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<document id="202906B87E0881A45B70187CE497EEC9" ID-DOI="10.1038/s42003-023-05210-5" ID-Zenodo-Dep="8279657" IM.bibliography_approvedBy="tatiana" IM.illustrations_approvedBy="tatiana" IM.materialsCitations_approvedBy="tatiana" IM.metadata_approvedBy="tatiana" IM.tables_approvedBy="tatiana" IM.taxonomicNames_approvedBy="tatiana" IM.treatments_approvedBy="tatiana" checkinTime="1692884671621" checkinUser="tatiana" docAuthor="Sevim-Erol, Ayla, Begun, D. R., Sözer, Ç. Sönmez, Mayda, S., van den Hoek Ostende, L. W., Martin, R. M. G. &amp; Alçiçek, M. Cihat" docDate="2023" docId="03D487E7FF9B0F5AFCC0FEB98E7CFE84" docLanguage="en" docName="CommunBiol.6.1-10.pdf" docOrigin="Communications Biology 6 (1), No. 842" docSource="http://dx.doi.org/10.1038/s42003-023-05210-5" docTitle="Anadoluvius Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek 2023, gen. nov." docType="treatment" docVersion="3" lastPageNumber="7" masterDocId="FFEDFF9FFF9A0F5CFF83FF898B6EFFAD" masterDocTitle="A new ape from Türkiye and the radiation of late Miocene hominines" masterLastPageNumber="10" masterPageNumber="1" pageNumber="2" updateTime="1692885179472" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
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<mods:title id="AF1D49C14C7D021A47FE776B5471ECE4">A new ape from Türkiye and the radiation of late Miocene hominines</mods:title>
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<mods:namePart id="316837122FC126C27E5957985A5DEB60">Begun, D. R.</mods:namePart>
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<mods:namePart id="3B2681AF0E5FA87862F28A16F44026EC">Mayda, S.</mods:namePart>
<mods:affiliation id="EBB625B772C6CC2C708DDF62D1E48A56">Ege University Fakulty of Science, Department of Biology, İzmir, Türkiye.</mods:affiliation>
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<mods:namePart id="021512ECEAFB3F9EF37FC895B3B6CFF3">van den Hoek Ostende, L. W.</mods:namePart>
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<mods:namePart id="88AEA7D0816AB8A26D9C2FA496709E5A">Martin, R. M. G.</mods:namePart>
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<mods:namePart id="62B632B09245F9194E782DECF5EEB478">Alçiçek, M. Cihat</mods:namePart>
<mods:affiliation id="0D0B296AC89EE0885AB5A4742D387ABF">Pamukkale University, Department of Geology, 20070 Denizli, Türkiye.</mods:affiliation>
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<taxonomicName id="4C7D4D72FF9B0F5DFCC0FEB988AFFEE8" authority="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek, 2023" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[835,961,304,325]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="1" pageNumber="2" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="B909EAE3FF9B0F5DFCC0FEB988AFFEE8" box="[835,961,304,325]" italics="true" pageId="1" pageNumber="2">Anadoluvius</emphasis>
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<taxonomicNameLabel id="A23A5798FF9B0F5DFC4AFEB98F49FEE8" box="[969,1063,304,325]" pageId="1" pageNumber="2" rank="genus">gen. nov.</taxonomicNameLabel>
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<paragraph id="8BC236F1FF9B0F5DFCA9FEE18EC4FEDB" blockId="1.[810,1450,355,382]" box="[810,1450,355,382]" pageId="1" pageNumber="2">
<emphasis id="B909EAE3FF9B0F5DFCA9FEE188FCFED0" box="[810,914,360,381]" italics="true" pageId="1" pageNumber="2">Synonomy</emphasis>
.
<taxonomicName id="4C7D4D72FF9B0F5DFC1CFEE18E61FED3" authority="Bonis and Melentis" authorityName="Bonis and Melentis" box="[927,1295,360,382]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9B0F5DFC1CFEE18F53FED0" box="[927,1085,360,381]" italics="true" pageId="1" pageNumber="2">Ouranopithecus</emphasis>
Bonis and Melentis
</taxonomicName>
: Güleç et al.
<bibRefCitation id="EFEC4B00FF9B0F5DFA14FEEA8EC4FEDB" author="Gulec, E. &amp; Sevim, A. &amp; Pehlevan, C. &amp; Kaya, F." box="[1431,1450,355,374]" journalOrPublisher="Anthropol. Sci." pageId="1" pageNumber="2" pagination="153 - 158" part="115" refId="ref10704" refString="28. Gulec, E., Sevim, A., Pehlevan, C. &amp; Kaya, F. A new great ape from the late Miocene of Turkey. Anthropol. Sci. 115, 153 - 158 (2007)." title="A new great ape from the late Miocene of Turkey" type="journal article" year="2007">28</bibRefCitation>
</paragraph>
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<emphasis id="B909EAE3FF9B0F5DFCA9FE2988C8FE18" box="[810,934,416,437]" italics="true" pageId="1" pageNumber="2">Type species</emphasis>
.
<taxonomicName id="4C7D4D72FF9B0F5DFC34FE168FEFFE19" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[951,1153,415,436]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="turkae" status="comb. nov.">
<emphasis id="B909EAE3FF9B0F5DFC34FE168FEFFE19" box="[951,1153,415,436]" italics="true" pageId="1" pageNumber="2">Anadoluvius turkae</emphasis>
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<taxonomicNameLabel id="A23A5798FF9B0F5DFB0EFE168E6CFE18" box="[1165,1282,415,437]" pageId="1" pageNumber="2" rank="species">comb. nov.</taxonomicNameLabel>
Sevim Erol et al. 2023.
</paragraph>
</subSubSection>
<subSubSection id="C367657AFF9B0F5DFCA9FE7A88FAFD88" pageId="1" pageNumber="2" type="etymology">
<paragraph id="8BC236F1FF9B0F5DFCA9FE7A88FAFD88" blockId="1.[810,1471,499,549]" pageId="1" pageNumber="2">
<emphasis id="B909EAE3FF9B0F5DFCA9FE7A88FFFDA5" box="[810,913,499,520]" italics="true" pageId="1" pageNumber="2">Etymology</emphasis>
. Anadolu is the modern Turkish word for Anatolia and Anatolian.
</paragraph>
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<materialsCitation id="3B153CACFF9B0F5DFCA9FDCE8F49FDDC" country="Türkiye" latitude="40.608887" location="Corakyerler" longLatPrecision="19" longitude="33.633614" municipality="Cankiri" pageId="1" pageNumber="2" specimenCount="1" specimenCount-male="1" stateProvince="Anatolian" typeStatus="holotype">
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<typeStatus id="54C68853FF9B0F5DFCA9FDCE88EDFDF1" box="[810,899,583,604]" pageId="1" pageNumber="2">Holotype</typeStatus>
</emphasis>
. CO-205, a fragmented but largely complete
<specimenCount id="9D7BFD78FF9B0F5DFACFFDCE8E12FDF0" box="[1356,1404,583,605]" pageId="1" pageNumber="2" type="male">male</specimenCount>
palate with LI1-M3 and RC-M2
</materialsCitation>
(Supplementary
<figureCitation id="13462A74FF9B0F5DFB5BFDEA8E75FDD5" box="[1240,1307,611,633]" captionStart="Fig" captionStartId="1.[810,840,1914,1933]" captionTargetBox="[822,1459,1560,1888]" captionTargetId="figure-852@1.[820,1460,1558,1890]" captionTargetPageId="1" captionText="Fig. 1 CO 2100/2800. A female partial cranium. From left to right, palatal, right lateral and anterior views." figureDoi="http://doi.org/10.5281/zenodo.8279659" httpUri="https://zenodo.org/record/8279659/files/figure.png" pageId="1" pageNumber="2">Figs. 1</figureCitation>
,
<figureCitation id="13462A74FF9B0F5DFAAAFDEA8E59FDD4" box="[1321,1335,611,633]" captionStart="Fig" captionStartId="2.[116,146,887,906]" captionTargetBox="[129,764,138,862]" captionTargetId="figure-874@2.[127,767,135,863]" captionTargetPageId="2" captionText="Fig. 2 Cross sectional anatomy of the palate in Anadoluvius and other hominids (not to scale). Ekembo and extant hominids redrawn from31 Rudapithecus modified from40. Ouranopithecus redrawn from41 based on a ct scan. The Ardipithecus specimen, modified from42, is a surface rendering derived from ct scans and does not show the cross section but the lateral aspect. The Ekembo specimen is based on BMNH 16664, the holotype of Ekembo nyanzae. The Rudapithecus specimens are RUD 12, a female, and RUD 44, a male. The photographs to the right of the line drawings of Rudapithecus are the original specimens. The Anadoluvius specimens are CO-2100/2800 (female, left) and CO-205 (male, right), with photographs of casts of the reconstructed specimens (see SI for details of the reconstruction.) Line drawings of Anadoluvius are original to this work." figureDoi="http://doi.org/10.5281/zenodo.8279661" httpUri="https://zenodo.org/record/8279661/files/figure.png" pageId="1" pageNumber="2">2</figureCitation>
).
</paragraph>
<paragraph id="8BC236F1FF9B0F5DFCA9FD128F5FFC91" blockId="1.[810,1472,666,995]" pageId="1" pageNumber="2">
<materialsCitation id="3B153CACFF9B0F5DFCA9FD128F55FD1D" box="[810,1083,667,690]" country="Türkiye" latitude="40.608887" location="Corakyerler" longLatPrecision="19" longitude="33.633614" municipality="Cankiri" pageId="1" pageNumber="2" specimenCount="1" stateProvince="Anatolian" typeStatus="paratype">
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<typeStatus id="54C68853FF9B0F5DFCA9FD1288E5FD1D" box="[810,907,667,688]" pageId="1" pageNumber="2">Paratypes</typeStatus>
</emphasis>
. CO-300 (RM
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);
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<materialsCitation id="3B153CACFF9B0F5DFBC5FD1388D9FD61" country="Türkiye" latitude="40.608887" location="Corakyerler" longLatPrecision="19" longitude="33.633614" municipality="Cankiri" pageId="1" pageNumber="2" specimenCount="1" specimenCount-male="1" stateProvince="Anatolian" typeStatus="paratype">
CO-305 (
<specimenCount id="9D7BFD78FF9B0F5DFB24FD138FB7FD1D" box="[1191,1241,666,688]" pageId="1" pageNumber="2" type="male">male</specimenCount>
mandibular fragment with RC-M
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);
</materialsCitation>
<materialsCitation id="3B153CACFF9B0F5DFC41FD3F8892FD45" country="Türkiye" latitude="40.608887" location="Corakyerler" longLatPrecision="19" longitude="33.633614" municipality="Cankiri" pageId="1" pageNumber="2" specimenCount="1" specimenCount-female="1" stateProvince="Anatolian" typeStatus="paratype">
CO-710 (
<specimenCount id="9D7BFD78FF9B0F5DFBA0FD3F8F09FD61" box="[1059,1127,694,716]" pageId="1" pageNumber="2" type="female">female</specimenCount>
mandibular fragment with RP
<subScript id="17F934B4FF9B0F5DFA2EFD498ED8FD63" attach="left" box="[1453,1462,704,718]" fontSize="6" pageId="1" pageNumber="2">3</subScript>
- M
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); CO-2100 (RI
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);
</materialsCitation>
<materialsCitation id="3B153CACFF9B0F5DFB87FD5B8F5FFC91" country="Türkiye" latitude="40.608887" location="Corakyerler" longLatPrecision="19" longitude="33.633614" municipality="Cankiri" pageId="1" pageNumber="2" specimenCount="1" specimenCount-female="1" stateProvince="Anatolian" typeStatus="paratype">
CO-2800 (female partial cranium with RC- M
<superScript id="7C089BB9FF9B0F5DFCC3FD608824FD51" attach="left" box="[832,842,745,764]" fontSize="8" pageId="1" pageNumber="2">2</superScript>
, portions of the right maxilla, maxillary frontal processes, frontal maxillary processes and most of the frontal bone) (
<figureCitation id="13462A74FF9B0F5DFAFCFC838ED4FCB2" box="[1407,1466,778,800]" captionStart="Fig" captionStartId="1.[810,840,1914,1933]" captionTargetBox="[822,1459,1560,1888]" captionTargetId="figure-852@1.[820,1460,1558,1890]" captionTargetPageId="1" captionText="Fig. 1 CO 2100/2800. A female partial cranium. From left to right, palatal, right lateral and anterior views." figureDoi="http://doi.org/10.5281/zenodo.8279659" httpUri="https://zenodo.org/record/8279659/files/figure.png" pageId="1" pageNumber="2">Fig. 1</figureCitation>
; Supplementary
<figureCitation id="13462A74FF9B0F5DFC48FCAF8F47FC91" box="[971,1065,805,828]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStartId-0="3.[116,146,1344,1363]" captionStartId-1="4.[116,146,733,752]" captionStartId-2="5.[116,146,766,785]" captionTargetBox-1="[156,1428,137,704]" captionTargetBox-2="[236,1352,137,739]" captionTargetId-0="figure-817@3.[127,767,135,1319]" captionTargetId-1="figure-626@4.[154,1434,135,708]" captionTargetId-2="figure-471@5.[234,1354,135,742]" captionTargetPageId-0="3" captionTargetPageId-1="4" captionTargetPageId-2="5" captionText-0="Fig. 3 3-D reconstruction of the left P3 to M1 of CO 300, showing the root, root canal and pulp chamber configurations. Supplementary Table 5 for a comparison of root formulae. Scale =10 mm." captionText-1="Fig. 4 Strict consensus cladograms. The four taxon sets each produced cladograms with the same topology whether character states were left unordered or a subset were ordered (see Methods and Supplementary Note 5 for details). a 18 OTUs. The four taxa with the fewest codable character states (Graecopithecus, 10%, Chororapithecus,13%, Samburupithecus, 18%, and Orrorin, 29%) were excluded, as was Sahelanthropus. Both Orrorin and Sahelanthropus were coded from published descriptions, which introduces uncertainty (DRB, who coded all characters in this analysis, was unable to code characters from these taxa through direct observation). b 19 OTUs, with Sahelanthropus added. c 20 OTUs with Orrorin. There is a decrease in resolution with the inclusion of Sahelanthropus and Orrorin but the tree topologies are otherwise consistent. Sahelantthropus is always recovered as a stem hominid and Orrorin as a hominin. The first three cladograms all recover a hominine clade that includes the thickly enameled Balkan taxa and the dryopithecins. d 23 OTUs, including all taxa. Little resolution remains among hominids, with recognized clades (pongines) unresolved. This cladogram also fails to recover Ouranopithecus as a hominine, which is otherwise a common result in previous analyses. Bremer support values, character states, character definitions and the character matrix (nexus) are all included in Supplementary Note 5 and Supplementary Data 3." captionText-2="Fig. 5 A phylogeny of the taxa included in this analysis consistent with most of the cladograms presented here. Taxa are positioned in chronological order without regard to geography,with most taxa only known from a limited time span. Exceptions are Ekembo and Sivapithecus, with longer time ranges,which are positioned roughly when they are most abundant, in both cases about mid-way in their known time ranges. The different colored “puddles” represent hominid clades and/or stages of evolution. These can also be imagined as pools of related species in somewhat delimited space and time with broad ancestordescendant relationships. The lines are disconnected to reflect the difficulty in identifying actual ancestor-descendant relationships, but that these relations can be estimated between “puddles”. Blue puddle taxa are stem hominids and are all confined to Africa. Among these taxa the relations of Samburupithecus and Chororapithecus are unresolved in the cladograms except in so far as they are excluded from the clade that includes all Eurasian taxa and crown hominins. Other lines of evidence suggest that these taxa are members of the early or middle Miocene radiation of early apes (see text). The orange puddles are the pongines, which probably have their origin within the middle Miocene puddle, although not necessarily any of the taxa included here (another taxon, Griphopithecus, known from Europe and Türkiye, would be a member of the blue puddle but was not included in the cladistic analyses).While both are pongines,Ankarapithecus lacks derived features shared by Sivapithecus and Pongo, so the line representing the relationship between the latter two bi-passes Ankarapithecus. The three green-shade puddles represent the hominine clades as defined here.Bright green are the dryopithecins,with the younger taxa Rudapithecus and Hispanopithecus depicted as closely related and descendant from any of the older taxa or an unknown taxon sharing attributes with these three. The light green puddle includes the Balkan and Anatolian taxa, likely to have descended from somewhere in the dryopithecin puddle.Possible ancestor-descendant relationships are depicted in this puddle. The darker green puddle represents the crown hominines. The various lineages diverge from unknown ancestors, but probably a member of either of the older green shade puddles.Gorillas diverge first, followed by chimpanzees and humans.Orrorin and Ardipithecus are depicted in a manner consistent with their sister clade status, without implying a direct ancestor-descendant relationship." figureDoi-0="http://doi.org/10.5281/zenodo.8279663" figureDoi-1="http://doi.org/10.5281/zenodo.8279665" figureDoi-2="http://doi.org/10.5281/zenodo.8279667" httpUri-0="https://zenodo.org/record/8279663/files/figure.png" httpUri-1="https://zenodo.org/record/8279665/files/figure.png" httpUri-2="https://zenodo.org/record/8279667/files/figure.png" pageId="1" pageNumber="2">Figs. 35</figureCitation>
)
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="C367657AFF9B0F5EFBB8FCAF89BDFA9F" lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2" type="discussion">
<paragraph id="8BC236F1FF9B0F5EFBB8FCAF89BDFA9F" blockId="1.[810,1472,666,995]" lastBlockId="2.[116,778,1252,1330]" lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2">
Detailed specimen descriptions and a revised diagnosis for this new taxon appear in the Supplementary Notes 2, 3. The hypodigm is curated in the Department of Anthropology, Ankara University. All samples used in this analysis are listed in Supplementary
<tableCitation id="C6FF034AFF9B0F5DFB00FC1C8FB4FC06" box="[1155,1242,917,939]" captionStart="Table 1" captionStartId="1.[127,184,1631,1653]" captionTargetBox="[127,767,1722,1956]" captionTargetId="graphics-713@1.[116,777,1609,1966]" captionText="Table 1 Hominine and pongine synapomorphies (18 taxa, unordered)." httpUri="http://table.plazi.org/id/DF026679FF9B0F5DFFFCF9D68B97F93F" pageId="1" pageNumber="2" tableUuid="DF026679FF9B0F5DFFFCF9D68B97F93F">Tables 1</tableCitation>
,
<tableCitation id="C6FF034AFF9B0F5DFB6AFC1F8F99FC06" box="[1257,1271,918,939]" captionStart="Table 2" captionStartId="1.[820,877,1053,1075]" captionTargetBox="[820,1460,1145,1482]" captionTargetId="graphics-777@1.[810,1472,1031,1491]" captionText="Table 2 Hominine and pongine synapomorphies (18 taxa, ordered)." httpUri="http://table.plazi.org/id/DF026679FF9B0F5DFCB7FB9488FBFBFD" pageId="1" pageNumber="2" tableUuid="DF026679FF9B0F5DFCB7FB9488FBFBFD">2</tableCitation>
. Measurements are provided in Supplementary Tables 3, 4. Supplementary Note 4 presents the results of a comprehensive quantitative analysis of the
<taxonomicName id="4C7D4D72FF980F5EFF27FB6D8A4CFB54" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[164,290,1252,1273]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFF27FB6D8A4CFB54" box="[164,290,1252,1273]" italics="true" pageId="2" pageNumber="3">Anadoluvius</emphasis>
</taxonomicName>
hypodigm. Supplementary Note 5 provides details of the phylogenetic analysis. Supplementary Note 6 provides historical, geological and biochronologic background.
</paragraph>
</subSubSection>
<caption id="DF026679FF9B0F5DFCB7FB9488FBFBFD" ID-Table-UUID="DF026679FF9B0F5DFCB7FB9488FBFBFD" httpUri="http://table.plazi.org/id/DF026679FF9B0F5DFCB7FB9488FBFBFD" pageId="1" pageNumber="2" startId="1.[820,877,1053,1075]" targetBox="[820,1460,1145,1482]" targetIsTable="true" targetPageId="1" targetType="table">
<paragraph id="8BC236F1FF9B0F5DFCB7FB9488FBFBFD" blockId="1.[820,1416,1053,1104]" pageId="1" pageNumber="2">Table 2 Hominine and pongine synapomorphies (18 taxa, ordered).</paragraph>
</caption>
<paragraph id="8BC236F1FF9B0F5DFCB7FBF08EDAFA65" pageId="1" pageNumber="2">
<table id="F97DC451FF9BF0A3FCB7FBF08EDAFA67" box="[820,1460,1145,1482]" gridcols="2" gridrows="13" pageId="1" pageNumber="2">
<tr id="354D34B3FF9BF0A3FCB7FBF08EDAFB21" box="[820,1460,1145,1164]" gridrow="0" pageId="1" pageNumber="2">
<th id="769C5DCFFF9BF0A3FCB7FBF08F47FB21" box="[820,1065,1145,1164]" gridcol="0" gridrow="0" pageId="1" pageNumber="2">Hominine</th>
<th id="769C5DCFFF9BF0A3FB2FFBF08EDAFB21" box="[1196,1460,1145,1164]" gridcol="1" gridrow="0" pageId="1" pageNumber="2">
<emphasis id="B909EAE3FF9B0F5DFB2FFBF08F9BFB21" bold="true" box="[1196,1269,1145,1164]" pageId="1" pageNumber="2">Pongine</emphasis>
</th>
</tr>
<tr id="354D34B3FF9BF0A3FCB7FB108EDAFB01" box="[820,1460,1177,1196]" gridrow="1" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FCB7FB108F47FB01" box="[820,1065,1177,1196]" gridcol="0" gridrow="1" pageId="1" pageNumber="2">Supraorbital torus</td>
<td id="769C5DCFFF9BF0A3FB2FFB108EDAFB01" box="[1196,1460,1177,1196]" gridcol="1" gridrow="1" pageId="1" pageNumber="2">Supraorbital margin</td>
</tr>
<tr id="354D34B3FF9BF0A3FCB7FB3A8EDAFB6B" box="[820,1460,1203,1222]" gridrow="2" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FCB7FB3A8F47FB6B" box="[820,1065,1203,1222]" gridcol="0" gridrow="2" pageId="1" pageNumber="2">Frontal sinus/glabella</td>
<td id="769C5DCFFF9BF0A3FB2FFB3A8EDAFB6B" box="[1196,1460,1203,1222]" gridcol="1" gridrow="2" pageId="1" pageNumber="2">Interorbital space</td>
</tr>
<tr id="354D34B3FF9BF0A3FCB7FB458EDAFB72" box="[820,1460,1228,1247]" gridrow="3" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FCB7FB458F47FB72" box="[820,1065,1228,1247]" gridcol="0" gridrow="3" pageId="1" pageNumber="2">Ethmoidal sinus</td>
<td id="769C5DCFFF9BF0A3FB2FFB458EDAFB72" box="[1196,1460,1228,1247]" gridcol="1" gridrow="3" pageId="1" pageNumber="2">Lateral orbital pillar surface</td>
</tr>
<tr id="354D34B3FF9BF0A3FCB7FB6A8EDAFB54" box="[820,1460,1251,1273]" gridrow="4" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FCB7FB6A8F47FB54" box="[820,1065,1251,1273]" gridcol="0" gridrow="4" pageId="1" pageNumber="2">
I
<superScript id="7C089BB9FF9B0F5DFCB9FB6A882EFB5F" attach="left" box="[826,832,1251,1266]" fontSize="6" pageId="1" pageNumber="2">1</superScript>
marginal ridge shape
</td>
<td id="769C5DCFFF9BF0A3FB2FFB6A8EDAFB54" box="[1196,1460,1251,1273]" gridcol="1" gridrow="4" pageId="1" pageNumber="2">Nasal bone length</td>
</tr>
<tr id="354D34B3FF9BF0A3FCB7FA898EDAFABE" box="[820,1460,1280,1299]" gridrow="5" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FCB7FA898F47FABE" box="[820,1065,1280,1299]" gridcol="0" gridrow="5" pageId="1" pageNumber="2">C implantation</td>
<td id="769C5DCFFF9BF0A3FB2FFA898EDAFABE" box="[1196,1460,1280,1299]" gridcol="1" gridrow="5" pageId="1" pageNumber="2">Nasal bone breath</td>
</tr>
<tr id="354D34B3FF9BF0A3FCB7FA938EDAFA80" box="[820,1460,1306,1325]" gridrow="6" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FCB7FA938F47FA80" box="[820,1065,1306,1325]" gridcol="0" gridrow="6" pageId="1" pageNumber="2">Upper premolar length</td>
<td id="769C5DCFFF9BF0A3FB2FFA938EDAFA80" box="[1196,1460,1306,1325]" gridcol="1" gridrow="6" pageId="1" pageNumber="2">Clivus length</td>
</tr>
<tr id="354D34B3FF9BF0A3FCB7FABD8EDAFAEA" box="[820,1460,1332,1351]" gridrow="7" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FCB7FABD8F47FAEA" box="[820,1065,1332,1351]" gridcol="0" gridrow="7" pageId="1" pageNumber="2">Lower molar buccal cristids</td>
<td id="769C5DCFFF9BF0A3FB2FFABD8EDAFAEA" box="[1196,1460,1332,1351]" gridcol="1" gridrow="7" pageId="1" pageNumber="2">Clivus orientation</td>
</tr>
<tr id="354D34B3FF9BF0A3FCB7FAC48EDAFACD" box="[820,1460,1357,1376]" gridrow="8" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FCB7FAC48F47FACD" box="[820,1065,1357,1376]" gridcol="0" gridrow="8" pageId="1" pageNumber="2">Supraorbital torus</td>
<td id="769C5DCFFF9BF0A3FB2FFAC48EDAFACD" box="[1196,1460,1357,1376]" gridcol="1" gridrow="8" pageId="1" pageNumber="2">C inclination</td>
</tr>
<tr id="354D34B3FF9BF0A3FCB7FAEE8EDAFAD7" box="[820,1460,1383,1402]" gridrow="9" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FCB7FAEE8EDAFAD7" box="[820,1460,1383,1402]" colspan="2" colspanRight="1" gridcol="0" gridrow="9" pageId="1" pageNumber="2">Incisive canal</td>
</tr>
<tr id="354D34B3FF9BF0A3FCB7FA088EDAFA39" box="[820,1460,1409,1428]" gridrow="10" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FCB7FA088EDAFA39" box="[820,1460,1409,1428]" colspan="2" colspanRight="1" gridcol="0" gridrow="10" pageId="1" pageNumber="2">Incisive foramen size</td>
</tr>
<tr id="354D34B3FF9BF0A3FCB7FA128EDAFA03" box="[820,1460,1435,1454]" gridrow="11" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FCB7FA128EDAFA03" box="[820,1460,1435,1454]" colspan="2" colspanRight="1" gridcol="0" gridrow="11" pageId="1" pageNumber="2">Upper premolar crests</td>
</tr>
<tr id="354D34B3FF9BF0A3FCB7FA3C8EDAFA67" box="[820,1460,1461,1482]" gridrow="12" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FCB7FA3C8EDAFA67" box="[820,1460,1461,1482]" colspan="2" colspanRight="1" gridcol="0" gridrow="12" pageId="1" pageNumber="2">
P
<subScript id="17F934B4FF9B0F5DFB3BFA348FAEFA67" attach="left" box="[1208,1216,1469,1482]" fontSize="5" pageId="1" pageNumber="2">3</subScript>
hypoproto-postprotocristid
</td>
</tr>
</table>
</paragraph>
<caption id="DF026679FF9B0F5DFFFCF9D68B97F93F" ID-Table-UUID="DF026679FF9B0F5DFFFCF9D68B97F93F" httpUri="http://table.plazi.org/id/DF026679FF9B0F5DFFFCF9D68B97F93F" pageId="1" pageNumber="2" startId="1.[127,184,1631,1653]" targetBox="[127,767,1722,1956]" targetIsTable="true" targetPageId="1" targetType="table">
<paragraph id="8BC236F1FF9B0F5DFFFCF9D68B97F93F" blockId="1.[127,719,1631,1682]" pageId="1" pageNumber="2">Table 1 Hominine and pongine synapomorphies (18 taxa, unordered).</paragraph>
</caption>
<paragraph id="8BC236F1FF9B0F5DFFFCF9338991F80F" pageId="1" pageNumber="2">
<table id="F97DC451FF9BF0A3FFFCF9338991F809" box="[127,767,1722,1956]" gridcols="2" gridrows="9" pageId="1" pageNumber="2">
<tr id="354D34B3FF9BF0A3FFFCF9338991F960" box="[127,767,1722,1741]" gridrow="0" pageId="1" pageNumber="2">
<th id="769C5DCFFF9BF0A3FFFCF9338A1AF960" box="[127,372,1722,1741]" gridcol="0" gridrow="0" pageId="1" pageNumber="2">Hominine</th>
<th id="769C5DCFFF9BF0A3FE74F9338991F960" box="[503,767,1722,1741]" gridcol="1" gridrow="0" pageId="1" pageNumber="2">
<emphasis id="B909EAE3FF9B0F5DFE74F933892EF960" bold="true" box="[503,576,1722,1741]" pageId="1" pageNumber="2">Pongine</emphasis>
</th>
</tr>
<tr id="354D34B3FF9BF0A3FFFCF9538991F940" box="[127,767,1754,1773]" gridrow="1" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FFFCF9538A1AF940" box="[127,372,1754,1773]" gridcol="0" gridrow="1" pageId="1" pageNumber="2">Supraorbital torus</td>
<td id="769C5DCFFF9BF0A3FE74F9538991F940" box="[503,767,1754,1773]" gridcol="1" gridrow="1" pageId="1" pageNumber="2">Supraorbital margin</td>
</tr>
<tr id="354D34B3FF9BF0A3FFFCF97D8991F8AA" box="[127,767,1780,1799]" gridrow="2" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FFFCF97D8A1AF8AA" box="[127,372,1780,1799]" gridcol="0" gridrow="2" pageId="1" pageNumber="2">Frontal sinus/glabella</td>
<td id="769C5DCFFF9BF0A3FE74F97D8991F8AA" box="[503,767,1780,1799]" gridcol="1" gridrow="2" pageId="1" pageNumber="2">Lateral orbital pillar surface</td>
</tr>
<tr id="354D34B3FF9BF0A3FFFCF8878991F88C" box="[127,767,1806,1825]" gridrow="3" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FFFCF8878A1AF88C" box="[127,372,1806,1825]" gridcol="0" gridrow="3" pageId="1" pageNumber="2">Ethmoidal sinus</td>
<td id="769C5DCFFF9BF0A3FE74F8878991F88C" box="[503,767,1806,1825]" gridcol="1" gridrow="3" pageId="1" pageNumber="2">Nasal bone length</td>
</tr>
<tr id="354D34B3FF9BF0A3FFFCF8AD8991F896" box="[127,767,1828,1851]" gridrow="4" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FFFCF8AD8A1AF896" box="[127,372,1828,1851]" gridcol="0" gridrow="4" pageId="1" pageNumber="2">
I
<superScript id="7C089BB9FF9B0F5DFF06F8AD8BE5F89E" attach="left" box="[133,139,1828,1843]" fontSize="6" pageId="1" pageNumber="2">1</superScript>
marginal ridge shape
</td>
<td id="769C5DCFFF9BF0A3FE74F8AD8991F896" box="[503,767,1828,1851]" gridcol="1" gridrow="4" pageId="1" pageNumber="2">Nasal bone breath</td>
</tr>
<tr id="354D34B3FF9BF0A3FFFCF8C88991F8F9" box="[127,767,1857,1876]" gridrow="5" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FFFCF8C88A1AF8F9" box="[127,372,1857,1876]" gridcol="0" gridrow="5" pageId="1" pageNumber="2">C implantation</td>
<td id="769C5DCFFF9BF0A3FE74F8C88991F8F9" box="[503,767,1857,1876]" gridcol="1" gridrow="5" pageId="1" pageNumber="2">Clivus orientation</td>
</tr>
<tr id="354D34B3FF9BF0A3FFFCF8D28991F8C3" box="[127,767,1883,1902]" gridrow="6" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FFFCF8D28A1AF8C3" box="[127,372,1883,1902]" gridcol="0" gridrow="6" pageId="1" pageNumber="2">Upper premolar length</td>
<td id="769C5DCFFF9BF0A3FE74F8D28991F8C3" box="[503,767,1883,1902]" gridcol="1" gridrow="6" pageId="1" pageNumber="2">C inclination</td>
</tr>
<tr id="354D34B3FF9BF0A3FFFCF8FC8991F825" box="[127,767,1909,1928]" gridrow="7" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FFFCF8FC8A1AF825" box="[127,372,1909,1928]" gridcol="0" gridrow="7" pageId="1" pageNumber="2">Lower molar buccal cristids</td>
<td id="769C5DCFFF9BF0A3FE74F8FC8991F825" box="[503,767,1909,1928]" gridcol="1" gridrow="7" pageId="1" pageNumber="2">Upper premolar crests</td>
</tr>
<tr id="354D34B3FF9BF0A3FFFCF8068991F809" box="[127,767,1935,1956]" gridrow="8" pageId="1" pageNumber="2">
<td id="769C5DCFFF9BF0A3FFFCF8068991F809" box="[127,767,1935,1956]" colspan="2" colspanRight="1" gridcol="0" gridrow="8" pageId="1" pageNumber="2">
P
<subScript id="17F934B4FF9B0F5DFD81F81E8964F809" attach="left" box="[514,522,1943,1956]" fontSize="5" pageId="1" pageNumber="2">3</subScript>
hypoproto-postprotocristid
</td>
</tr>
</table>
</paragraph>
<caption id="DF026679FF9B0F5DFCA9F8F38F2CF804" ID-DOI="http://doi.org/10.5281/zenodo.8279659" ID-Zenodo-Dep="8279659" httpUri="https://zenodo.org/record/8279659/files/figure.png" pageId="1" pageNumber="2" startId="1.[810,840,1914,1933]" targetBox="[822,1459,1560,1888]" targetPageId="1" targetType="figure">
<paragraph id="8BC236F1FF9B0F5DFCA9F8F38F2CF804" blockId="1.[810,1473,1914,1961]" pageId="1" pageNumber="2">
<emphasis id="B909EAE3FF9B0F5DFCA9F8F3889DF820" bold="true" box="[810,1011,1914,1933]" pageId="1" pageNumber="2">Fig. 1 CO 2100/2800.</emphasis>
A female partial cranium. From left to right, palatal, right lateral and anterior views.
</paragraph>
</caption>
<caption id="DF026679FF980F5EFFF7FCFE8984FB10" ID-DOI="http://doi.org/10.5281/zenodo.8279661" ID-Zenodo-Dep="8279661" httpUri="https://zenodo.org/record/8279661/files/figure.png" pageId="2" pageNumber="3" startId="2.[116,146,887,906]" targetBox="[129,764,138,862]" targetPageId="2" targetType="figure">
<paragraph id="8BC236F1FF980F5EFFF7FCFE8984FB10" blockId="2.[116,778,887,1214]" pageId="2" pageNumber="3">
<emphasis id="B909EAE3FF980F5EFFF7FCFE8A3CFC0B" bold="true" pageId="2" pageNumber="3">
Fig. 2 Cross sectional anatomy of the palate in
<taxonomicName id="4C7D4D72FF980F5EFDAFFCFE89F6FC26" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[556,664,887,907]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFDAFFCFE89F6FC26" bold="true" box="[556,664,887,907]" italics="true" pageId="2" pageNumber="3">Anadoluvius</emphasis>
</taxonomicName>
and other hominids (not to scale).
</emphasis>
<taxonomicName id="4C7D4D72FF980F5EFEDBFC1A8AF6FC0A" authorityName="McNulty, Begun, Kelley, Manthi &amp; Mbua" authorityYear="2015" box="[344,408,915,935]" class="Mammalia" genus="Ekembo" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFEDBFC1A8AF6FC0A" box="[344,408,915,935]" italics="true" pageId="2" pageNumber="3">Ekembo</emphasis>
</taxonomicName>
and extant hominids redrawn from
<superScript id="7C089BB9FF980F5EFD55FC19898BFC32" attach="left" box="[726,741,912,927]" fontSize="6" pageId="2" pageNumber="3">31</superScript>
<taxonomicName id="4C7D4D72FF980F5EFFF7FC278B8FFC6F" authorityName="Kretzoi" authorityYear="1969" box="[116,225,942,962]" class="Mammalia" family="Hominidae" genus="Rudapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFFF7FC278B8FFC6F" box="[116,225,942,962]" italics="true" pageId="2" pageNumber="3">Rudapithecus</emphasis>
</taxonomicName>
modified from
<superScript id="7C089BB9FF980F5EFEE0FC258A19FC16" attach="left" box="[355,375,940,955]" fontSize="6" pageId="2" pageNumber="3">40</superScript>
.
<taxonomicName id="4C7D4D72FF980F5EFE02FC27896CFC6F" box="[385,514,942,962]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFE02FC27896CFC6F" box="[385,514,942,962]" italics="true" pageId="2" pageNumber="3">Ouranopithecus</emphasis>
</taxonomicName>
redrawn from
<bibRefCitation id="EFEC4B00FF980F5EFDFCFC2589E1FC16" author="Bonis, Lde &amp; Melentis, J." box="[639,655,940,955]" journalOrPublisher="C. R. Acad. Sci. Paris" pageId="2" pageNumber="3" pagination="767 - 769" part="304" refId="ref11238" refString="41. Bonis, Lde &amp; Melentis, J. Interet de l' anatomie naso-maxillaire pour la phylogenie de Hominidae. C. R. Acad. Sci. Paris 304, 767 - 769 (1987)." title="Interet de l' anatomie naso-maxillaire pour la phylogenie de Hominidae" type="journal article" year="1987">
<superScript id="7C089BB9FF980F5EFDFCFC2589E1FC16" attach="left" box="[639,655,940,955]" fontSize="6" pageId="2" pageNumber="3">41</superScript>
</bibRefCitation>
based on a ct scan. The
<taxonomicName id="4C7D4D72FF980F5EFF52FC438A56FC73" box="[209,312,970,990]" class="Mammalia" family="Hominidae" genus="Ardipithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFF52FC438A56FC73" box="[209,312,970,990]" italics="true" pageId="2" pageNumber="3">Ardipithecus</emphasis>
</taxonomicName>
specimen, modified from
<bibRefCitation id="EFEC4B00FF980F5EFD98FC4E8940FC7B" author="White, T. D. &amp; et al." box="[539,558,967,982]" journalOrPublisher="Science" pageId="2" pageNumber="3" pagination="75 - 86" part="64" refId="ref11277" refString="42. White, T. D. et al. Ardipithecus ramidus and the paleobiology of early Hominids. Science 326, 64, 75 - 86 (2009)." title="Ardipithecus ramidus and the paleobiology of early Hominids" type="journal article" year="2009">
<superScript id="7C089BB9FF980F5EFD98FC4E8940FC7B" attach="left" box="[539,558,967,982]" fontSize="6" pageId="2" pageNumber="3">42</superScript>
</bibRefCitation>
, is a surface rendering derived from ct scans and does not show the cross section but the lateral aspect. The
<taxonomicName id="4C7D4D72FF980F5EFF61FB8B8A4CFBBB" authorityName="McNulty, Begun, Kelley, Manthi &amp; Mbua" authorityYear="2015" box="[226,290,1026,1046]" class="Mammalia" genus="Ekembo" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFF61FB8B8A4CFBBB" box="[226,290,1026,1046]" italics="true" pageId="2" pageNumber="3">Ekembo</emphasis>
</taxonomicName>
specimen is based on BMNH 16664, the holotype of
<taxonomicName id="4C7D4D72FF980F5EFFF7FB978A6EFB9F" baseAuthorityName="Le Gros Clark &amp; Leakey" baseAuthorityYear="1950" box="[116,256,1054,1074]" class="Mammalia" genus="Ekembo" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="nyanzae">
<emphasis id="B909EAE3FF980F5EFFF7FB978A6EFB9F" box="[116,256,1054,1074]" italics="true" pageId="2" pageNumber="3">Ekembo nyanzae</emphasis>
</taxonomicName>
. The
<taxonomicName id="4C7D4D72FF980F5EFEB7FB978ACFFB9F" authorityName="Kretzoi" authorityYear="1969" box="[308,417,1054,1074]" class="Mammalia" family="Hominidae" genus="Rudapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFEB7FB978ACFFB9F" box="[308,417,1054,1074]" italics="true" pageId="2" pageNumber="3">Rudapithecus</emphasis>
</taxonomicName>
specimens are RUD 12, a female, and RUD 44, a male. The photographs to the right of the line drawings of
<taxonomicName id="4C7D4D72FF980F5EFFF7FBDF8B8FFBC7" authorityName="Kretzoi" authorityYear="1969" box="[116,225,1110,1130]" class="Mammalia" family="Hominidae" genus="Rudapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFFF7FBDF8B8FFBC7" box="[116,225,1110,1130]" italics="true" pageId="2" pageNumber="3">Rudapithecus</emphasis>
</taxonomicName>
are the original specimens. The
<taxonomicName id="4C7D4D72FF980F5EFD8BFBDF8903FBC7" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[520,621,1110,1130]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFD8BFBDF8903FBC7" box="[520,621,1110,1130]" italics="true" pageId="2" pageNumber="3">Anadoluvius</emphasis>
</taxonomicName>
specimens are CO-2100/2800 (female, left) and CO-205 (male, right), with photographs of casts of the reconstructed specimens (see SI for details of the reconstruction.) Line drawings of
<taxonomicName id="4C7D4D72FF980F5EFE21FB238969FB13" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[418,519,1194,1214]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFE21FB238969FB13" box="[418,519,1194,1214]" italics="true" pageId="2" pageNumber="3">Anadoluvius</emphasis>
</taxonomicName>
are original to this work.
</paragraph>
</caption>
<subSubSection id="C367657AFF980F59FFF7FADD88ADF8CD" lastPageId="5" lastPageNumber="6" pageId="2" pageNumber="3" type="description">
<paragraph id="8BC236F1FF980F5EFFF7FADD8980F915" blockId="2.[116,778,1364,1972]" pageId="2" pageNumber="3">
<emphasis id="B909EAE3FF980F5EFFF7FADD8BA1FAC4" box="[116,207,1364,1385]" italics="true" pageId="2" pageNumber="3">Cranium</emphasis>
. The CO-2100/2800 partial cranium (
<figureCitation id="13462A74FF980F5EFDEFFADD89C5FAC4" box="[620,683,1364,1386]" captionStart="Fig" captionStartId="1.[810,840,1914,1933]" captionTargetBox="[822,1459,1560,1888]" captionTargetId="figure-852@1.[820,1460,1558,1890]" captionTargetPageId="1" captionText="Fig. 1 CO 2100/2800. A female partial cranium. From left to right, palatal, right lateral and anterior views." figureDoi="http://doi.org/10.5281/zenodo.8279659" httpUri="https://zenodo.org/record/8279659/files/figure.png" pageId="2" pageNumber="3">Fig. 1</figureCitation>
, Supplementary
<figureCitation id="13462A74FF980F5EFF51FAF98A78FA28" box="[210,278,1392,1414]" captionStart="Fig" captionStartId="2.[116,146,887,906]" captionTargetBox="[129,764,138,862]" captionTargetId="figure-874@2.[127,767,135,863]" captionTargetPageId="2" captionText="Fig. 2 Cross sectional anatomy of the palate in Anadoluvius and other hominids (not to scale). Ekembo and extant hominids redrawn from31 Rudapithecus modified from40. Ouranopithecus redrawn from41 based on a ct scan. The Ardipithecus specimen, modified from42, is a surface rendering derived from ct scans and does not show the cross section but the lateral aspect. The Ekembo specimen is based on BMNH 16664, the holotype of Ekembo nyanzae. The Rudapithecus specimens are RUD 12, a female, and RUD 44, a male. The photographs to the right of the line drawings of Rudapithecus are the original specimens. The Anadoluvius specimens are CO-2100/2800 (female, left) and CO-205 (male, right), with photographs of casts of the reconstructed specimens (see SI for details of the reconstruction.) Line drawings of Anadoluvius are original to this work." figureDoi="http://doi.org/10.5281/zenodo.8279661" httpUri="https://zenodo.org/record/8279661/files/figure.png" pageId="2" pageNumber="3">Figs. 2</figureCitation>
,
<figureCitation id="13462A74FF980F5EFEA6FAF98A5CFA2B" box="[293,306,1392,1414]" captionStart="Fig" captionStartId="3.[116,146,1344,1363]" captionTargetId="figure-817@3.[127,767,135,1319]" captionTargetPageId="3" captionText="Fig. 3 3-D reconstruction of the left P3 to M1 of CO 300, showing the root, root canal and pulp chamber configurations. Supplementary Table 5 for a comparison of root formulae. Scale =10 mm." figureDoi="http://doi.org/10.5281/zenodo.8279663" httpUri="https://zenodo.org/record/8279663/files/figure.png" pageId="2" pageNumber="3">3</figureCitation>
) was recovered with some crushing and displacement of several broken pieces (see SM for restoration details and detailed description.) The frontal bone is nearly intact, missing only portions within the temporal fossa and the squama approaching bregma. This distinguishes it from the most complete facial specimen of
<taxonomicName id="4C7D4D72FF980F5EFEB5FA728ABDF9BD" box="[310,467,1531,1552]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFEB5FA728ABDF9BD" box="[310,467,1531,1552]" italics="true" pageId="2" pageNumber="3">Ouranopithecus</emphasis>
</taxonomicName>
(XIR-1), which is broken just beyond the superior orbital margins and preserves almost nothing of the frontal squama (Supplementary
<figureCitation id="13462A74FF980F5EFD81F9BA8950F9E4" box="[514,574,1587,1609]" captionStart="Fig" captionStartId="2.[116,146,887,906]" captionTargetBox="[129,764,138,862]" captionTargetId="figure-874@2.[127,767,135,863]" captionTargetPageId="2" captionText="Fig. 2 Cross sectional anatomy of the palate in Anadoluvius and other hominids (not to scale). Ekembo and extant hominids redrawn from31 Rudapithecus modified from40. Ouranopithecus redrawn from41 based on a ct scan. The Ardipithecus specimen, modified from42, is a surface rendering derived from ct scans and does not show the cross section but the lateral aspect. The Ekembo specimen is based on BMNH 16664, the holotype of Ekembo nyanzae. The Rudapithecus specimens are RUD 12, a female, and RUD 44, a male. The photographs to the right of the line drawings of Rudapithecus are the original specimens. The Anadoluvius specimens are CO-2100/2800 (female, left) and CO-205 (male, right), with photographs of casts of the reconstructed specimens (see SI for details of the reconstruction.) Line drawings of Anadoluvius are original to this work." figureDoi="http://doi.org/10.5281/zenodo.8279661" httpUri="https://zenodo.org/record/8279661/files/figure.png" pageId="2" pageNumber="3">Fig. 2</figureCitation>
). Though damaged, the position and orientation of the premaxilla is better preserved in CO-2100/2800 than in CO-205 and
<taxonomicName id="4C7D4D72FF980F5EFE6CF9E289E2F92D" box="[495,652,1643,1664]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFE6CF9E289E2F92D" box="[495,652,1643,1664]" italics="true" pageId="2" pageNumber="3">Ouranopithecus</emphasis>
</taxonomicName>
(XIR-1 and RPl 128), confirming previous interpretations of a stepped and overlapping morphology in these specimens
<bibRefCitation id="EFEC4B00FF980F5EFDACF9178957F91C" author="Begun, D. R." box="[559,569,1694,1713]" editor="Henke, W. &amp; Tattersall, I." journalOrPublisher="Springer" pageId="2" pageNumber="3" pagination="1261 - 1332" refId="ref9660" refString="1. Begun, D. R. Fossil Record of Miocene Hominoids in Handbook of Paleoanthropology (eds. Henke, W. &amp; Tattersall, I.) 1261 - 1332 (Springer, 2015)." title="Fossil Record of Miocene Hominoids" type="book chapter" volumeTitle="Handbook of Paleoanthropology" year="2015">1</bibRefCitation>
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,
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,
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(
<figureCitation id="13462A74FF980F5EFD25F92B898EF915" box="[678,736,1698,1720]" captionStart="Fig" captionStartId="2.[116,146,887,906]" captionTargetBox="[129,764,138,862]" captionTargetId="figure-874@2.[127,767,135,863]" captionTargetPageId="2" captionText="Fig. 2 Cross sectional anatomy of the palate in Anadoluvius and other hominids (not to scale). Ekembo and extant hominids redrawn from31 Rudapithecus modified from40. Ouranopithecus redrawn from41 based on a ct scan. The Ardipithecus specimen, modified from42, is a surface rendering derived from ct scans and does not show the cross section but the lateral aspect. The Ekembo specimen is based on BMNH 16664, the holotype of Ekembo nyanzae. The Rudapithecus specimens are RUD 12, a female, and RUD 44, a male. The photographs to the right of the line drawings of Rudapithecus are the original specimens. The Anadoluvius specimens are CO-2100/2800 (female, left) and CO-205 (male, right), with photographs of casts of the reconstructed specimens (see SI for details of the reconstruction.) Line drawings of Anadoluvius are original to this work." figureDoi="http://doi.org/10.5281/zenodo.8279661" httpUri="https://zenodo.org/record/8279661/files/figure.png" pageId="2" pageNumber="3">Fig. 2</figureCitation>
).
</paragraph>
<paragraph id="8BC236F1FF980F5EFF0DF937889DFECC" blockId="2.[116,778,1364,1972]" lastBlockId="2.[810,1472,136,354]" pageId="2" pageNumber="3">
The premaxilla of
<taxonomicName id="4C7D4D72FF980F5EFECFF9378AA4F97E" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[332,458,1726,1747]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFECFF9378AA4F97E" box="[332,458,1726,1747]" italics="true" pageId="2" pageNumber="3">Anadoluvius</emphasis>
</taxonomicName>
is short and vertical compared with
<taxonomicName id="4C7D4D72FF980F5EFF2AF9528BBFF95D" authorityName="Oken" authorityYear="1816" box="[169,209,1755,1776]" class="Mammalia" family="Hominidae" genus="Pan" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFF2AF9528BBFF95D" box="[169,209,1755,1776]" italics="true" pageId="2" pageNumber="3">Pan</emphasis>
</taxonomicName>
,
<taxonomicName id="4C7D4D72FF980F5EFF5DF9528A75F95D" authorityName="Lacepede" authorityYear="1799" box="[222,283,1755,1776]" class="Mammalia" family="Hominidae" genus="Pongo" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFF5DF9528A75F95D" box="[222,283,1755,1776]" italics="true" pageId="2" pageNumber="3">Pongo</emphasis>
</taxonomicName>
, and australopithecines, and is most like
<emphasis id="B909EAE3FF980F5EFD47F9538867F942" box="[708,777,1754,1775]" italics="true" pageId="2" pageNumber="3">Gorilla</emphasis>
and dryopithecins, being relatively short in the alveolar portion but expanded nasally to overlap with the palatine process of the maxilla (
<figureCitation id="13462A74FF980F5EFF48F8A78A6AF8EE" box="[203,260,1838,1860]" captionStart="Fig" captionStartId="2.[116,146,887,906]" captionTargetBox="[129,764,138,862]" captionTargetId="figure-874@2.[127,767,135,863]" captionTargetPageId="2" captionText="Fig. 2 Cross sectional anatomy of the palate in Anadoluvius and other hominids (not to scale). Ekembo and extant hominids redrawn from31 Rudapithecus modified from40. Ouranopithecus redrawn from41 based on a ct scan. The Ardipithecus specimen, modified from42, is a surface rendering derived from ct scans and does not show the cross section but the lateral aspect. The Ekembo specimen is based on BMNH 16664, the holotype of Ekembo nyanzae. The Rudapithecus specimens are RUD 12, a female, and RUD 44, a male. The photographs to the right of the line drawings of Rudapithecus are the original specimens. The Anadoluvius specimens are CO-2100/2800 (female, left) and CO-205 (male, right), with photographs of casts of the reconstructed specimens (see SI for details of the reconstruction.) Line drawings of Anadoluvius are original to this work." figureDoi="http://doi.org/10.5281/zenodo.8279661" httpUri="https://zenodo.org/record/8279661/files/figure.png" pageId="2" pageNumber="3">Fig. 2</figureCitation>
) The incisor alveoli are positioned along the mesial transverse plane of the canine crowns (
<figureCitation id="13462A74FF980F5EFD85F8C38924F8F2" box="[518,586,1866,1888]" captionStart="Fig" captionStartId="1.[810,840,1914,1933]" captionTargetBox="[822,1459,1560,1888]" captionTargetId="figure-852@1.[820,1460,1558,1890]" captionTargetPageId="1" captionText="Fig. 1 CO 2100/2800. A female partial cranium. From left to right, palatal, right lateral and anterior views." figureDoi="http://doi.org/10.5281/zenodo.8279659" httpUri="https://zenodo.org/record/8279659/files/figure.png" pageId="2" pageNumber="3">Figs. 1</figureCitation>
,
<figureCitation id="13462A74FF980F5EFDDBF8C3890AF8F2" box="[600,612,1866,1887]" captionStart="Fig" captionStartId="2.[116,146,887,906]" captionTargetBox="[129,764,138,862]" captionTargetId="figure-874@2.[127,767,135,863]" captionTargetPageId="2" captionText="Fig. 2 Cross sectional anatomy of the palate in Anadoluvius and other hominids (not to scale). Ekembo and extant hominids redrawn from31 Rudapithecus modified from40. Ouranopithecus redrawn from41 based on a ct scan. The Ardipithecus specimen, modified from42, is a surface rendering derived from ct scans and does not show the cross section but the lateral aspect. The Ekembo specimen is based on BMNH 16664, the holotype of Ekembo nyanzae. The Rudapithecus specimens are RUD 12, a female, and RUD 44, a male. The photographs to the right of the line drawings of Rudapithecus are the original specimens. The Anadoluvius specimens are CO-2100/2800 (female, left) and CO-205 (male, right), with photographs of casts of the reconstructed specimens (see SI for details of the reconstruction.) Line drawings of Anadoluvius are original to this work." figureDoi="http://doi.org/10.5281/zenodo.8279661" httpUri="https://zenodo.org/record/8279661/files/figure.png" pageId="2" pageNumber="3">2</figureCitation>
; Supplementary
<figureCitation id="13462A74FF980F5EFFF7F8EF8BC0F8D1" box="[116,174,1894,1916]" captionStart="Fig" captionStartId="5.[116,146,766,785]" captionTargetBox="[236,1352,137,739]" captionTargetId="figure-471@5.[234,1354,135,742]" captionTargetPageId="5" captionText="Fig. 5 A phylogeny of the taxa included in this analysis consistent with most of the cladograms presented here. Taxa are positioned in chronological order without regard to geography,with most taxa only known from a limited time span. Exceptions are Ekembo and Sivapithecus, with longer time ranges,which are positioned roughly when they are most abundant, in both cases about mid-way in their known time ranges. The different colored “puddles” represent hominid clades and/or stages of evolution. These can also be imagined as pools of related species in somewhat delimited space and time with broad ancestordescendant relationships. The lines are disconnected to reflect the difficulty in identifying actual ancestor-descendant relationships, but that these relations can be estimated between “puddles”. Blue puddle taxa are stem hominids and are all confined to Africa. Among these taxa the relations of Samburupithecus and Chororapithecus are unresolved in the cladograms except in so far as they are excluded from the clade that includes all Eurasian taxa and crown hominins. Other lines of evidence suggest that these taxa are members of the early or middle Miocene radiation of early apes (see text). The orange puddles are the pongines, which probably have their origin within the middle Miocene puddle, although not necessarily any of the taxa included here (another taxon, Griphopithecus, known from Europe and Türkiye, would be a member of the blue puddle but was not included in the cladistic analyses).While both are pongines,Ankarapithecus lacks derived features shared by Sivapithecus and Pongo, so the line representing the relationship between the latter two bi-passes Ankarapithecus. The three green-shade puddles represent the hominine clades as defined here.Bright green are the dryopithecins,with the younger taxa Rudapithecus and Hispanopithecus depicted as closely related and descendant from any of the older taxa or an unknown taxon sharing attributes with these three. The light green puddle includes the Balkan and Anatolian taxa, likely to have descended from somewhere in the dryopithecin puddle.Possible ancestor-descendant relationships are depicted in this puddle. The darker green puddle represents the crown hominines. The various lineages diverge from unknown ancestors, but probably a member of either of the older green shade puddles.Gorillas diverge first, followed by chimpanzees and humans.Orrorin and Ardipithecus are depicted in a manner consistent with their sister clade status, without implying a direct ancestor-descendant relationship." figureDoi="http://doi.org/10.5281/zenodo.8279667" httpUri="https://zenodo.org/record/8279667/files/figure.png" pageId="2" pageNumber="3">Fig. 5</figureCitation>
) In two specimens of
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<emphasis id="B909EAE3FF980F5EFE11F8EF895EF8D6" box="[402,560,1894,1915]" italics="true" pageId="2" pageNumber="3">Ouranopithecus</emphasis>
</taxonomicName>
(RPl 128 and XIR 1) the upper incisors are well anterior to the canines (Supplementary
<figureCitation id="13462A74FF980F5EFFF7F8178BDEF81E" box="[116,176,1949,1972]" captionStart="Fig" captionStartId="5.[116,146,766,785]" captionTargetBox="[236,1352,137,739]" captionTargetId="figure-471@5.[234,1354,135,742]" captionTargetPageId="5" captionText="Fig. 5 A phylogeny of the taxa included in this analysis consistent with most of the cladograms presented here. Taxa are positioned in chronological order without regard to geography,with most taxa only known from a limited time span. Exceptions are Ekembo and Sivapithecus, with longer time ranges,which are positioned roughly when they are most abundant, in both cases about mid-way in their known time ranges. The different colored “puddles” represent hominid clades and/or stages of evolution. These can also be imagined as pools of related species in somewhat delimited space and time with broad ancestordescendant relationships. The lines are disconnected to reflect the difficulty in identifying actual ancestor-descendant relationships, but that these relations can be estimated between “puddles”. Blue puddle taxa are stem hominids and are all confined to Africa. Among these taxa the relations of Samburupithecus and Chororapithecus are unresolved in the cladograms except in so far as they are excluded from the clade that includes all Eurasian taxa and crown hominins. Other lines of evidence suggest that these taxa are members of the early or middle Miocene radiation of early apes (see text). The orange puddles are the pongines, which probably have their origin within the middle Miocene puddle, although not necessarily any of the taxa included here (another taxon, Griphopithecus, known from Europe and Türkiye, would be a member of the blue puddle but was not included in the cladistic analyses).While both are pongines,Ankarapithecus lacks derived features shared by Sivapithecus and Pongo, so the line representing the relationship between the latter two bi-passes Ankarapithecus. The three green-shade puddles represent the hominine clades as defined here.Bright green are the dryopithecins,with the younger taxa Rudapithecus and Hispanopithecus depicted as closely related and descendant from any of the older taxa or an unknown taxon sharing attributes with these three. The light green puddle includes the Balkan and Anatolian taxa, likely to have descended from somewhere in the dryopithecin puddle.Possible ancestor-descendant relationships are depicted in this puddle. The darker green puddle represents the crown hominines. The various lineages diverge from unknown ancestors, but probably a member of either of the older green shade puddles.Gorillas diverge first, followed by chimpanzees and humans.Orrorin and Ardipithecus are depicted in a manner consistent with their sister clade status, without implying a direct ancestor-descendant relationship." figureDoi="http://doi.org/10.5281/zenodo.8279667" httpUri="https://zenodo.org/record/8279667/files/figure.png" pageId="2" pageNumber="3">Fig. 5</figureCitation>
) In NKT 89 the premaxilla is severely damaged, but the posterior edge of the lateral incisor appears to be aligned with the anterior transverse plane of the canines, a position most like
<taxonomicName id="4C7D4D72FF980F5EFCA9FF4988C6FF78" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[810,936,192,213]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFCA9FF4988C6FF78" box="[810,936,192,213]" italics="true" pageId="2" pageNumber="3">Anadoluvius</emphasis>
</taxonomicName>
. The frontal bone of
<taxonomicName id="4C7D4D72FF980F5EFB1AFF498E79FF78" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[1177,1303,192,213]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFB1AFF498E79FF78" box="[1177,1303,192,213]" italics="true" pageId="2" pageNumber="3">Anadoluvius</emphasis>
</taxonomicName>
differs strongly from that of
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<emphasis id="B909EAE3FF980F5EFC32FF558F21FF5C" box="[945,1103,220,241]" italics="true" pageId="2" pageNumber="3">Ouranopithecus</emphasis>
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in the smooth biconvex squama of the former, contrasting with a broad concavity above glabella in the latter. The superior orbital margins of
<taxonomicName id="4C7D4D72FF980F5EFB75FE9D8EFAFE84" box="[1270,1428,276,297]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFB75FE9D8EFAFE84" box="[1270,1428,276,297]" italics="true" pageId="2" pageNumber="3">Ouranopithecus</emphasis>
</taxonomicName>
are broad, rounded and slightly projecting while they are sharp and flat in
<taxonomicName id="4C7D4D72FF980F5EFCECFEC58883FECC" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[879,1005,332,353]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFCECFEC58883FECC" box="[879,1005,332,353]" italics="true" pageId="2" pageNumber="3">Anadoluvius</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BC236F1FF980F5EFCA9FE0A8FA6FDF0" blockId="2.[810,1472,387,1274]" pageId="2" pageNumber="3">
<emphasis id="B909EAE3FF980F5EFCA9FE0A88E7FE35" box="[810,905,387,408]" italics="true" pageId="2" pageNumber="3">Mandible</emphasis>
. A principal components analysis based on mandibular measurements available for
<emphasis id="B909EAE3FF980F5EFBCFFE168EE1FE19" box="[1100,1423,415,436]" italics="true" pageId="2" pageNumber="3">
<taxonomicName id="4C7D4D72FF980F5EFBCFFE168F8BFE19" authorityName="von Königswald" authorityYear="1972" box="[1100,1253,415,436]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">Graecopithecus</taxonomicName>
,
<taxonomicName id="4C7D4D72FF980F5EFB72FE168EE1FE19" box="[1265,1423,415,436]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">Ouranopithecus</taxonomicName>
</emphasis>
and
<taxonomicName id="4C7D4D72FF980F5EFCA9FE3288C6FE7D" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[810,936,443,464]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFCA9FE3288C6FE7D" box="[810,936,443,464]" italics="true" pageId="2" pageNumber="3">Anadoluvius</emphasis>
</taxonomicName>
is presented in Supplementary Fig. 6 and Supplementary Data 1). The Çorakyerler, Nikiti 1 and
<taxonomicName id="4C7D4D72FF980F5EFAAAFE5E8EAEFE41" authorityName="von Königswald" authorityYear="1972" box="[1321,1472,471,492]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFAAAFE5E8EAEFE41" box="[1321,1472,471,492]" italics="true" pageId="2" pageNumber="3">Graecopithecus</emphasis>
</taxonomicName>
mandibles are separated from each other, especially along PC 2, and from
<taxonomicName id="4C7D4D72FF980F5EFC18FD868F57FD89" box="[923,1081,527,548]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFC18FD868F57FD89" box="[923,1081,527,548]" italics="true" pageId="2" pageNumber="3">Ouranopithecus</emphasis>
</taxonomicName>
, illustrating the diversity present in these samples. Supplementary Data 1 includes the data matrix, summary statistics, scores, and loadings.
</paragraph>
<paragraph id="8BC236F1FF980F5EFCC0FDEA8E01FC91" blockId="2.[810,1472,387,1274]" pageId="2" pageNumber="3">
<taxonomicName id="4C7D4D72FF980F5EFCC0FDEA88D2FDD5" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[835,956,611,632]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFCC0FDEA88D2FDD5" box="[835,956,611,632]" italics="true" pageId="2" pageNumber="3">Anadoluvius</emphasis>
</taxonomicName>
, like
<taxonomicName id="4C7D4D72FF980F5EFC74FDEA8FE6FDD5" authorityName="von Königswald" authorityYear="1972" box="[1015,1160,611,632]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFC74FDEA8FE6FDD5" box="[1015,1160,611,632]" italics="true" pageId="2" pageNumber="3">Graecopithecus</emphasis>
</taxonomicName>
, and NKT 21, has a relatively narrow mandible compared with the combined sex sample of
<taxonomicName id="4C7D4D72FF980F5EFCA9FD1388ADFD02" box="[810,963,666,687]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFCA9FD1388ADFD02" box="[810,963,666,687]" italics="true" pageId="2" pageNumber="3">Ouranopithecus</emphasis>
</taxonomicName>
(Supplementary Fig. 7a). Supplementary Fig. 7b compares relative mandibular corpus breadth at each tooth position (P
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-M
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) in
<emphasis id="B909EAE3FF980F5EFC33FD5B8FBDFD4A" box="[944,1235,722,743]" italics="true" pageId="2" pageNumber="3">
<taxonomicName id="4C7D4D72FF980F5EFC33FD5B8F42FD4A" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[944,1068,722,743]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">Anadoluvius</taxonomicName>
,
<taxonomicName id="4C7D4D72FF980F5EFBC1FD5B8FBDFD4A" authorityName="von Königswald" authorityYear="1972" box="[1090,1235,722,743]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">Graecopithecus</taxonomicName>
</emphasis>
, and
<taxonomicName id="4C7D4D72FF980F5EFAA2FD5B8ED7FD4A" box="[1313,1465,722,743]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFAA2FD5B8ED7FD4A" box="[1313,1465,722,743]" italics="true" pageId="2" pageNumber="3">Ouranopithecus</emphasis>
</taxonomicName>
.
<taxonomicName id="4C7D4D72FF980F5EFCA9FD6788CAFCAE" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[810,932,750,771]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFCA9FD6788CAFCAE" box="[810,932,750,771]" italics="true" pageId="2" pageNumber="3">Anadoluvius</emphasis>
</taxonomicName>
is similar in mandibular robusticity at the premolar level but at the level of the molars it matches or strongly exceeds the maximum value in the other taxa (Supplementary Fig. 7b).
</paragraph>
<paragraph id="8BC236F1FF980F5EFCC0FCCB8FBBFB57" blockId="2.[810,1472,387,1274]" pageId="2" pageNumber="3">
There is diversity in mandibular robusticity (breadth relative to height) and in dental size ratios as well among the samples of eastern Mediterranean apes (Supplementary Fig. 8ae).
<taxonomicName id="4C7D4D72FF980F5EFAF7FCF3880BFC07" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFAF7FCF3880BFC07" italics="true" pageId="2" pageNumber="3">Anadoluvius</emphasis>
</taxonomicName>
is distinct from
<taxonomicName id="4C7D4D72FF980F5EFBAAFC1C8FD1FC07" authorityName="von Königswald" authorityYear="1972" box="[1065,1215,917,938]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFBAAFC1C8FD1FC07" box="[1065,1215,917,938]" italics="true" pageId="2" pageNumber="3">Graecopithecus</emphasis>
</taxonomicName>
in all mandibular and dental ratios.
<taxonomicName id="4C7D4D72FF980F5EFC34FC388F5BFC6B" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[951,1077,945,966]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFC34FC388F5BFC6B" box="[951,1077,945,966]" italics="true" pageId="2" pageNumber="3">Anadoluvius</emphasis>
</taxonomicName>
falls beyond the range of variation of
<taxonomicName id="4C7D4D72FF980F5EFCA9FC4488A6FC4F" box="[810,968,973,994]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFCA9FC4488A6FC4F" box="[810,968,973,994]" italics="true" pageId="2" pageNumber="3">Ouranopithecus</emphasis>
</taxonomicName>
in relative corpus breadth at M
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-M
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(Supplementary Fig. 8a) and M
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size (Supplementary Fig. 8d). Interestingly, NKT 21 falls outside the
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<emphasis id="B909EAE3FF980F5EFBF2FB8C8E61FBB7" box="[1137,1295,1029,1050]" italics="true" pageId="2" pageNumber="3">Ouranopithecus</emphasis>
</taxonomicName>
range in relative P
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length and M
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size (Supplementary Fig. 8df). It has a relatively short symphyseal-molar distance, at the 25% quartile for the Ravin sample, and a relatively robust mandible at M
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-M
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, at the 75% quartile for
<taxonomicName id="4C7D4D72FF980F5EFBA1FBFC8FAEFB27" box="[1058,1216,1141,1162]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFBA1FBFC8FAEFB27" box="[1058,1216,1141,1162]" italics="true" pageId="2" pageNumber="3">Ouranopithecus</emphasis>
</taxonomicName>
(Supplementary Fig. 8a, c). In
<taxonomicName id="4C7D4D72FF980F5EFCEFFB188F6CFB0B" authorityName="von Königswald" authorityYear="1972" box="[876,1026,1169,1190]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFCEFFB188F6CFB0B" box="[876,1026,1169,1190]" italics="true" pageId="2" pageNumber="3">Graecopithecus</emphasis>
</taxonomicName>
the symphysis is positioned closest to the molars, just barely in the range of the Ravin sample. In summary, in most quantitative comparisons
<taxonomicName id="4C7D4D72FF980F5EFB16FB418E7DFB70" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[1173,1299,1224,1245]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFB16FB418E7DFB70" box="[1173,1299,1224,1245]" italics="true" pageId="2" pageNumber="3">Anadoluvius</emphasis>
</taxonomicName>
is distinguished from
<taxonomicName id="4C7D4D72FF980F5EFCE7FB6D8F6CFB54" box="[868,1026,1252,1273]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFCE7FB6D8F6CFB54" box="[868,1026,1252,1273]" italics="true" pageId="2" pageNumber="3">Ouranopithecus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF980F5EFBBAFB6D8FA1FB54" authorityName="von Königswald" authorityYear="1972" box="[1081,1231,1252,1273]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFBBAFB6D8FA1FB54" box="[1081,1231,1252,1273]" italics="true" pageId="2" pageNumber="3">Graecopithecus</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BC236F1FF980F5EFCA9FA958FF0FA10" blockId="2.[810,1472,1308,1748]" pageId="2" pageNumber="3">
<emphasis id="B909EAE3FF980F5EFCA9FA958FEAFA9C" box="[810,1156,1308,1330]" italics="true" pageId="2" pageNumber="3">Tooth roots and enamel thickness</emphasis>
.
<figureCitation id="13462A74FF980F5EFB17FA958F84FA9C" box="[1172,1258,1308,1330]" captionStart="Fig" captionStartId="3.[116,146,1344,1363]" captionTargetId="figure-817@3.[127,767,135,1319]" captionTargetPageId="3" captionText="Fig. 3 3-D reconstruction of the left P3 to M1 of CO 300, showing the root, root canal and pulp chamber configurations. Supplementary Table 5 for a comparison of root formulae. Scale =10 mm." figureDoi="http://doi.org/10.5281/zenodo.8279663" httpUri="https://zenodo.org/record/8279663/files/figure.png" pageId="2" pageNumber="3">Figure 3</figureCitation>
shows the root and root canal morphology of CO-300, the male mandible of
<taxonomicName id="4C7D4D72FF980F5EFA0FFAB18810FAC4" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFA0FFAB18810FAC4" italics="true" pageId="2" pageNumber="3">Anadoluvius</emphasis>
</taxonomicName>
(see Methods for segmentation details). Unlike
<taxonomicName id="4C7D4D72FF980F5EFA0CFADD88CEFA28" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFA0CFADD88CEFA28" italics="true" pageId="2" pageNumber="3">Ouranopithecus</emphasis>
</taxonomicName>
, the distal roots of P
<subScript id="17F934B4FF980F5EFB0EFAF08FF8FA2A" attach="left" box="[1165,1174,1401,1415]" fontSize="6" pageId="2" pageNumber="3">3</subScript>
to M
<subScript id="17F934B4FF980F5EFB5AFAF08F8CFA2A" attach="left" box="[1241,1250,1401,1415]" fontSize="6" pageId="2" pageNumber="3">1</subScript>
in
<taxonomicName id="4C7D4D72FF980F5EFA8CFAF98EE3FA28" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[1295,1421,1392,1413]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFA8CFAF98EE3FA28" box="[1295,1421,1392,1413]" italics="true" pageId="2" pageNumber="3">Anadoluvius</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF980F5EFCA9FA0588AEFA0C" authorityName="von Königswald" authorityYear="1972" box="[810,960,1420,1441]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFCA9FA0588AEFA0C" box="[810,960,1420,1441]" italics="true" pageId="2" pageNumber="3">Graecopithecus</emphasis>
</taxonomicName>
<bibRefCitation id="EFEC4B00FF980F5EFC43FA0E88BDFA37" author="Fuss, J. &amp; Spassov, N. &amp; Begun, D. R. &amp; Bohme, M." box="[960,979,1415,1434]" journalOrPublisher="PLoS ONE" pageId="2" pageNumber="3" pagination="0177127" part="12" refId="ref10011" refString="11. Fuss, J., Spassov, N., Begun, D. R. &amp; Bohme, M. Potential hominin affinities of Graecopithecus from the Late Miocene of Europe. PLoS ONE 12, e 0177127 (2017)." title="Potential hominin affinities of Graecopithecus from the Late Miocene of Europe" type="journal article" year="2017">
<superScript id="7C089BB9FF980F5EFC43FA0E88BDFA37" attach="left" box="[960,979,1415,1434]" fontSize="8" pageId="2" pageNumber="3">11</superScript>
</bibRefCitation>
are single fused roots with two root canals (
<figureCitation id="13462A74FF980F5EFCB2FA218802FA10" box="[817,876,1448,1470]" captionStart="Fig" captionStartId="3.[116,146,1344,1363]" captionTargetId="figure-817@3.[127,767,135,1319]" captionTargetPageId="3" captionText="Fig. 3 3-D reconstruction of the left P3 to M1 of CO 300, showing the root, root canal and pulp chamber configurations. Supplementary Table 5 for a comparison of root formulae. Scale =10 mm." figureDoi="http://doi.org/10.5281/zenodo.8279663" httpUri="https://zenodo.org/record/8279663/files/figure.png" pageId="2" pageNumber="3">Fig. 3</figureCitation>
and Supplementary Table 5).
</paragraph>
<paragraph id="8BC236F1FF980F5EFCC0FA4A8F02F979" blockId="2.[810,1472,1308,1748]" pageId="2" pageNumber="3">
Like
<emphasis id="B909EAE3FF980F5EFCFBFA4A8FCBFA75" box="[888,1189,1475,1496]" italics="true" pageId="2" pageNumber="3">
<taxonomicName id="4C7D4D72FF980F5EFCFBFA4A8F76FA75" box="[888,1048,1475,1496]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">Ouranopithecus</taxonomicName>
,
<taxonomicName id="4C7D4D72FF980F5EFBA4FA4A8FCBFA75" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[1063,1189,1475,1496]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">Anadoluvius</taxonomicName>
</emphasis>
has thick enamel. Supplementary Fig. 9 illustrates ranges of variation in relative enamel thickness (RET) in the M
<subScript id="17F934B4FF980F5EFBABF98C8F5FF9BE" attach="left" box="[1064,1073,1541,1555]" fontSize="6" pageId="2" pageNumber="3">2</subScript>
of Miocene, Plio-Pleistocene and living hominoids.
<taxonomicName id="4C7D4D72FF980F5EFC27F99E8F4CF981" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[932,1058,1559,1580]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFC27F99E8F4CF981" box="[932,1058,1559,1580]" italics="true" pageId="2" pageNumber="3">Anadoluvius</emphasis>
</taxonomicName>
has thicker enamel than most Miocene apes, falling at the upper end of the range in
<taxonomicName id="4C7D4D72FF980F5EFA8FF9BA8EE7F9E5" authorityName="Leakey &amp; Leakey" authorityYear="1986" box="[1292,1417,1587,1608]" class="Mammalia" family="Victoriapithecidae" genus="Afropithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFA8FF9BA8EE7F9E5" box="[1292,1417,1587,1608]" italics="true" pageId="2" pageNumber="3">Afropithecus</emphasis>
</taxonomicName>
<bibRefCitation id="EFEC4B00FF980F5EFA0AF9A78EF2F9EC" author="Smith, T. M. &amp; Tafforeau, P. &amp; Pouech, J. &amp; Begun, D. R." box="[1417,1436,1582,1601]" journalOrPublisher="J. Hum. Evol." pageId="2" pageNumber="3" pagination="102649" part="136" refId="ref10785" refString="30. Smith, T. M., Tafforeau, P., Pouech, J. &amp; Begun, D. R. Enamel thickness and dental development in Rudapithecus hungaricus. J. Hum. Evol. 136, 102649 (2019)." title="Enamel thickness and dental development in Rudapithecus hungaricus" type="journal article" year="2019">
<superScript id="7C089BB9FF980F5EFA0AF9A78EF2F9EC" attach="left" box="[1417,1436,1582,1601]" fontSize="8" pageId="2" pageNumber="3">30</superScript>
</bibRefCitation>
Its RET is greater than RPl 641, an M
<subScript id="17F934B4FF980F5EFB33F9D08FD7F9CA" attach="left" box="[1200,1209,1625,1639]" fontSize="6" pageId="2" pageNumber="3">3</subScript>
of
<taxonomicName id="4C7D4D72FF980F5EFB64F9C68EEBF9C9" box="[1255,1413,1615,1636]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFB64F9C68EEBF9C9" box="[1255,1413,1615,1636]" italics="true" pageId="2" pageNumber="3">Ouranopithecus</emphasis>
</taxonomicName>
. The relationship between M
<subScript id="17F934B4FF980F5EFB9AF9FD8F4CF92F" attach="left" box="[1049,1058,1652,1666]" fontSize="6" pageId="2" pageNumber="3">2</subScript>
and M
<subScript id="17F934B4FF980F5EFBEEF9FD8F18F92F" attach="left" box="[1133,1142,1652,1666]" fontSize="6" pageId="2" pageNumber="3">3</subScript>
RET is variable in hominoids
<bibRefCitation id="EFEC4B00FF980F5EFA25F9EF8ED7F9D4" author="Smith, T. M. &amp; Tafforeau, P. &amp; Pouech, J. &amp; Begun, D. R." box="[1446,1465,1638,1657]" journalOrPublisher="J. Hum. Evol." pageId="2" pageNumber="3" pagination="102649" part="136" refId="ref10785" refString="30. Smith, T. M., Tafforeau, P., Pouech, J. &amp; Begun, D. R. Enamel thickness and dental development in Rudapithecus hungaricus. J. Hum. Evol. 136, 102649 (2019)." title="Enamel thickness and dental development in Rudapithecus hungaricus" type="journal article" year="2019">
<superScript id="7C089BB9FF980F5EFA25F9EF8ED7F9D4" attach="left" box="[1446,1465,1638,1657]" fontSize="8" pageId="2" pageNumber="3">30</superScript>
</bibRefCitation>
, but their ranges of variation always overlap.
<taxonomicName id="4C7D4D72FF980F5EFB66F90E8E0DF931" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[1253,1379,1671,1692]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFB66F90E8E0DF931" box="[1253,1379,1671,1692]" italics="true" pageId="2" pageNumber="3">Anadoluvius</emphasis>
</taxonomicName>
falls well above the ranges in extant hominids and within the
<taxonomicName id="4C7D4D72FF980F5EFACAF92A8ED1F915" box="[1353,1471,1699,1720]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="afarensis">
<emphasis id="B909EAE3FF980F5EFACAF92A8ED1F915" box="[1353,1471,1699,1720]" italics="true" pageId="2" pageNumber="3">A. afarensis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF980F5EFCDBF93688BDF979" box="[856,979,1727,1748]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="africanus">
<emphasis id="B909EAE3FF980F5EFCDBF93688BDF979" box="[856,979,1727,1748]" italics="true" pageId="2" pageNumber="3">A. africanus</emphasis>
</taxonomicName>
75% quartiles.
</paragraph>
<paragraph id="8BC236F1FF980F5EFCA9F97E8E52F819" blockId="2.[810,1472,1782,1972]" pageId="2" pageNumber="3">
<emphasis id="B909EAE3FF980F5EFCA9F97E88F0F8A1" box="[810,926,1783,1804]" italics="true" pageId="2" pageNumber="3">Canine size</emphasis>
. The results of an ANOVA examining lower canine relative size is presented in Supplementary Table 7. In canine size relative to the geometric mean
<taxonomicName id="4C7D4D72FF980F5EFBFCF8A78E73F8EE" box="[1151,1309,1838,1859]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFBFCF8A78E73F8EE" box="[1151,1309,1838,1859]" italics="true" pageId="2" pageNumber="3">Ouranopithecus</emphasis>
</taxonomicName>
is significantly different from extant African apes in having relatively small canines. The relative size of the CO-305 mandibular canine (0.53) is equal to the mean of
<taxonomicName id="4C7D4D72FF980F5EFB99F80B8FD6F83A" box="[1050,1208,1922,1943]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFB99F80B8FD6F83A" box="[1050,1208,1922,1943]" italics="true" pageId="2" pageNumber="3">Ouranopithecus</emphasis>
</taxonomicName>
males and at the low end of the range of variation in
<taxonomicName id="4C7D4D72FF980F5EFBC9F8178F1CF81E" authorityName="Oken" authorityYear="1816" box="[1098,1138,1950,1971]" class="Mammalia" family="Hominidae" genus="Pan" kingdom="Animalia" order="Primates" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF980F5EFBC9F8178F1CF81E" box="[1098,1138,1950,1971]" italics="true" pageId="2" pageNumber="3">Pan</emphasis>
</taxonomicName>
males and females.
</paragraph>
<caption id="DF026679FF990F5FFFF7FAC98959FA27" ID-DOI="http://doi.org/10.5281/zenodo.8279663" ID-Zenodo-Dep="8279663" httpUri="https://zenodo.org/record/8279663/files/figure.png" pageId="3" pageNumber="4" startId="3.[116,146,1344,1363]" targetBox="[129,765,138,1317]" targetPageId="3" targetType="figure">
<paragraph id="8BC236F1FF990F5FFFF7FAC98959FA27" blockId="3.[116,777,1344,1419]" pageId="3" pageNumber="4">
<emphasis id="B909EAE3FF990F5FFFF7FAC98957FAC2" bold="true" pageId="3" pageNumber="4">Fig. 3 3-D reconstruction of the left P3 to M1 of CO 300, showing the root, root canal and pulp chamber configurations.</emphasis>
Supplementary Table 5 for a comparison of root formulae. Scale =10 mm.
</paragraph>
</caption>
<paragraph id="8BC236F1FF990F5FFF0DFA4B8A93F8F3" blockId="3.[116,778,1474,1970]" pageId="3" pageNumber="4">
Canines are small in the Balkan/Anatolian sample compared with other fossil and extant apes including
<taxonomicName id="4C7D4D72FF990F5FFDC6FA5789ADFA5E" box="[581,707,1502,1523]" class="Mammalia" family="Hominidae" genus="Ardipithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFDC6FA5789ADFA5E" box="[581,707,1502,1523]" italics="true" pageId="3" pageNumber="4">Ardipithecus</emphasis>
</taxonomicName>
, being more consistent with the ranges in
<taxonomicName id="4C7D4D72FF990F5FFE5EFA7389EDF9A2" authorityName="R.A.Dart" authorityYear="1925" box="[477,643,1530,1551]" class="Mammalia" family="Hominidae" genus="Australopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFE5EFA7389EDF9A2" box="[477,643,1530,1551]" italics="true" pageId="3" pageNumber="4">Australopithecus</emphasis>
</taxonomicName>
. The ratio of lower canine to M
<subScript id="17F934B4FF990F5FFEADF9968A59F980" attach="left" box="[302,311,1567,1581]" fontSize="6" pageId="3" pageNumber="4">2</subScript>
size (canine maximum ln x bd/M
<superScript id="7C089BB9FF990F5FFD12F99889F5F989" attach="left" box="[657,667,1553,1572]" fontSize="8" pageId="3" pageNumber="4">2</superScript>
maximum ln x bd) in
<taxonomicName id="4C7D4D72FF990F5FFF7AF9B88A19F9EB" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[249,375,1585,1606]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFF7AF9B88A19F9EB" box="[249,375,1585,1606]" italics="true" pageId="3" pageNumber="4">Anadoluvius</emphasis>
</taxonomicName>
is lower than in any male and most females except
<taxonomicName id="4C7D4D72FF990F5FFE94F9C48AD3F9CF" authorityName="R.A.Dart" authorityYear="1925" box="[279,445,1613,1634]" class="Mammalia" family="Hominidae" genus="Australopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFE94F9C48AD3F9CF" box="[279,445,1613,1634]" italics="true" pageId="3" pageNumber="4">Australopithecus</emphasis>
</taxonomicName>
(Supplementary Fig. 10a). The Balkan/Anatolian specimens have relatively small canines compared with African apes when scaled to the individual geometric means (Supplementary Fig. 10b and Supplementary Tables 6, 7; geometric mean of 12 variables). Scaled M
<subScript id="17F934B4FF990F5FFDC0F94E8922F978" attach="left" box="[579,588,1735,1749]" fontSize="6" pageId="3" pageNumber="4">2</subScript>
size is large and beyond the range of variation of African apes in
<emphasis id="B909EAE3FF990F5FFDEEF9508B96F8A7" italics="true" pageId="3" pageNumber="4">
<taxonomicName id="4C7D4D72FF990F5FFDEEF9508868F943" authorityName="von Königswald" authorityYear="1972" box="[621,774,1753,1774]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Graecopithecus</taxonomicName>
,
<taxonomicName id="4C7D4D72FF990F5FFFF7F97C8B9AF8A7" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[116,244,1781,1802]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Anadoluvius</taxonomicName>
,
</emphasis>
and Nikiti 1while Ravin de la Pluie
<taxonomicName id="4C7D4D72FF990F5FFDE8F97C8867F8A7" box="[619,777,1781,1802]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFDE8F97C8867F8A7" box="[619,777,1781,1802]" italics="true" pageId="3" pageNumber="4">Ouranopithecus</emphasis>
</taxonomicName>
is intermediate between the other fossil apes and
<emphasis id="B909EAE3FF990F5FFD47F8988867F88B" box="[708,777,1809,1830]" italics="true" pageId="3" pageNumber="4">Gorilla</emphasis>
(Supplementary Fig. 10c). Canine size is compared to each tooth position in Supplementary Fig. 11a, b.
</paragraph>
<paragraph id="8BC236F1FF990F5FFF0DF8ED8802FD1D" blockId="3.[116,778,1474,1970]" lastBlockId="3.[810,1472,135,1972]" pageId="3" pageNumber="4">
The results of a cladistic analysis using a data matrix of 112 characters and between 18 and 23 taxa are presented in
<figureCitation id="13462A74FF990F5FFD49F809886AF838" box="[714,772,1920,1942]" captionStart="Fig" captionStartId="4.[116,146,733,752]" captionTargetBox="[156,1428,137,704]" captionTargetId="figure-626@4.[154,1434,135,708]" captionTargetPageId="4" captionText="Fig. 4 Strict consensus cladograms. The four taxon sets each produced cladograms with the same topology whether character states were left unordered or a subset were ordered (see Methods and Supplementary Note 5 for details). a 18 OTUs. The four taxa with the fewest codable character states (Graecopithecus, 10%, Chororapithecus,13%, Samburupithecus, 18%, and Orrorin, 29%) were excluded, as was Sahelanthropus. Both Orrorin and Sahelanthropus were coded from published descriptions, which introduces uncertainty (DRB, who coded all characters in this analysis, was unable to code characters from these taxa through direct observation). b 19 OTUs, with Sahelanthropus added. c 20 OTUs with Orrorin. There is a decrease in resolution with the inclusion of Sahelanthropus and Orrorin but the tree topologies are otherwise consistent. Sahelantthropus is always recovered as a stem hominid and Orrorin as a hominin. The first three cladograms all recover a hominine clade that includes the thickly enameled Balkan taxa and the dryopithecins. d 23 OTUs, including all taxa. Little resolution remains among hominids, with recognized clades (pongines) unresolved. This cladogram also fails to recover Ouranopithecus as a hominine, which is otherwise a common result in previous analyses. Bremer support values, character states, character definitions and the character matrix (nexus) are all included in Supplementary Note 5 and Supplementary Data 3." figureDoi="http://doi.org/10.5281/zenodo.8279665" httpUri="https://zenodo.org/record/8279665/files/figure.png" pageId="3" pageNumber="4">Fig. 4</figureCitation>
,
<tableCitation id="C6FF034AFF990F5FFFF7F8158BA5F81C" box="[116,203,1948,1970]" captionStart="Table 1" captionStartId="1.[127,184,1631,1653]" captionTargetBox="[127,767,1722,1956]" captionTargetId="graphics-713@1.[116,777,1609,1966]" captionText="Table 1 Hominine and pongine synapomorphies (18 taxa, unordered)." httpUri="http://table.plazi.org/id/DF026679FF9B0F5DFFFCF9D68B97F93F" pageId="3" pageNumber="4" tableUuid="DF026679FF9B0F5DFFFCF9D68B97F93F">Tables 1</tableCitation>
,
<tableCitation id="C6FF034AFF990F5FFF59F8158B88F81C" box="[218,230,1948,1969]" captionStart="Table 2" captionStartId="1.[820,877,1053,1075]" captionTargetBox="[820,1460,1145,1482]" captionTargetId="graphics-777@1.[810,1472,1031,1491]" captionText="Table 2 Hominine and pongine synapomorphies (18 taxa, ordered)." httpUri="http://table.plazi.org/id/DF026679FF9B0F5DFCB7FB9488FBFBFD" pageId="3" pageNumber="4" tableUuid="DF026679FF9B0F5DFCB7FB9488FBFBFD">2</tableCitation>
and in the supplementary materials (Supplementary Figs. 1214; Supplementary Data 2, 3; Supplementary Note 5). All four cladograms are strict consensus. The analysis was run both with all characters unordered and 21 of the 112 characters ordered (see Methods and Supplementary Note 5 for character matrix assumptions). Both analyses yield similar results. The tree topologies are identical, and predictably the tree values are lower in the analyses using ordered characters. All but the cladograms that include the taxa with more than 80% missing data recover a clade that includes Eurasian apes and hominines to the exclusion of pongines and stem hominids (Fig. 4). The potential problem of using data from published sources as opposed to direct observation is illustrated in the results for
<taxonomicName id="4C7D4D72FF990F5FFB6AFE328EEDFE7D" box="[1257,1411,443,464]" class="Mammalia" family="Hominidae" genus="Sahelanthropus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFB6AFE328EEDFE7D" box="[1257,1411,443,464]" italics="true" pageId="3" pageNumber="4">Sahelanthropus</emphasis>
</taxonomicName>
. This taxon, universally attributed to the
<taxonomicName id="4C7D4D72FF990F5FFB4DFE5E8E2EFE40" authorityName="Gray" authorityYear="1825" box="[1230,1344,471,493]" class="Mammalia" family="Hominidae" kingdom="Animalia" order="Primates" pageId="3" pageNumber="2" phylum="Chordata" rank="subFamily" subFamily="Homininae">Homininae</taxonomicName>
and most commonly to the Hominini, is never recovered in these positions in these analyses, being consistently a stem hominid.
<taxonomicName id="4C7D4D72FF990F5FFAEBFD86881BFDED" class="Mammalia" family="Hominidae" genus="Sahelanthropus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFAEBFD86881BFDED" italics="true" pageId="3" pageNumber="4">Sahelanthropus</emphasis>
</taxonomicName>
could be coded for 72% of the characters. The potential for coding inconsistent with the criteria used to code other taxa is therefore larger than for
<taxonomicName id="4C7D4D72FF990F5FFBB3FDEA8F13FDD5" authorityName="Senut, Pickford, Gommery, Mein, Cheboi &amp; Coppens" authorityYear="2001" box="[1072,1149,611,632]" class="Mammalia" family="Hominidae" genus="Orrorin" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFBB3FDEA8F13FDD5" box="[1072,1149,611,632]" italics="true" pageId="3" pageNumber="4">Orrorin</emphasis>
</taxonomicName>
, which is also coded from the literature, but for which only 29% of the characters could be coded.
</paragraph>
<paragraph id="8BC236F1FF990F5FFCC0FD3E8F17FCFA" blockId="3.[810,1472,135,1972]" pageId="3" pageNumber="4">We mapped synapomorphies and Bremer support values onto two consensus cladograms (Supplementary Figs. 12, 13). Table 1 lists the hominine synapomorphies of the cladogram (ordered and unordered) with the fewest missing data (18 OTUs). A phylogeny consistent with a large majority of the cladograms presented here appears in Fig. 5.</paragraph>
<paragraph id="8BC236F1FF990F5FFCC0FCD788E9FAC7" blockId="3.[810,1472,135,1972]" pageId="3" pageNumber="4">
Supplementary Fig. 14 shows the results of the analyses of all four taxon sets with all character states ordered. Unlike the unordered and partly ordered analyses, the cladograms with differing OTUs vary widely. The 19 OTU (including
<taxonomicName id="4C7D4D72FF990F5FFAEBFC38881BFC4F" class="Mammalia" family="Hominidae" genus="Sahelanthropus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFAEBFC38881BFC4F" italics="true" pageId="3" pageNumber="4">Sahelanthropus</emphasis>
</taxonomicName>
) fully ordered analysis is consistent with previous ones while the 18 and 20 OTU analyses result in a pongine clade including all European and Anatolian taxa, which contrasts with all previous cladistic results from analyses with large data sets and numerical cladistic methods (e.g. TNT, PAUP, etc). The 20 OTU analysis includes several other unconventional results such as an
<emphasis id="B909EAE3FF990F5FFCA9FBFC8890FB27" box="[810,1022,1141,1162]" italics="true" pageId="3" pageNumber="4">
<taxonomicName id="4C7D4D72FF990F5FFCA9FBFC8819FB27" authorityName="Senut, Pickford, Gommery, Mein, Cheboi &amp; Coppens" authorityYear="2001" box="[810,887,1141,1162]" class="Mammalia" family="Hominidae" genus="Orrorin" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Orrorin</taxonomicName>
-
<taxonomicName id="4C7D4D72FF990F5FFC03FBFC8890FB27" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[896,1022,1141,1162]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Anadoluvius</taxonomicName>
</emphasis>
clade that, along with
<taxonomicName id="4C7D4D72FF990F5FFB62FBFC8E11FB27" box="[1249,1407,1141,1162]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFB62FBFC8E11FB27" box="[1249,1407,1141,1162]" italics="true" pageId="3" pageNumber="4">Ouranopithecus</emphasis>
</taxonomicName>
as the sister to pongines, and a pongine clade that includes all European and Anatolian taxa. The 23 OTU analysis remains highly unresolved. Given the diversity of results of these all ordered analyses and the broader consistency of the analyses in which the data matrices were either unordered or partly ordered, we consider the latter to be more reliable (see Methods for a discussion of our rationale on ordering character states in this analysis).
</paragraph>
<paragraph id="8BC236F1FF990F5FFCC0FAF98E1EF979" blockId="3.[810,1472,135,1972]" pageId="3" pageNumber="4">
The consensus cladogram with the three taxa represented by &lt;20% of the character matrix (23 OTUs) is uninformative, given the low level of resolution. The cladograms resulting from the unordered or partly ordered analyses, which exclude these taxa (
<taxonomicName id="4C7D4D72FF990F5FFCB1FA5688B8FA59" authorityName="Suwa, Kono, Katoh, Asfaw &amp; Beyene" authorityYear="2007" box="[818,982,1503,1524]" class="Mammalia" genus="Chororapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFCB1FA5688B8FA59" box="[818,982,1503,1524]" italics="true" pageId="3" pageNumber="4">Chororapithecus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C7D4D72FF990F5FFC71FA568FF0FA59" authorityName="Ishida &amp; Pickford" authorityYear="1997" box="[1010,1182,1503,1524]" class="Mammalia" genus="Samburupithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFC71FA568FF0FA59" box="[1010,1182,1503,1524]" italics="true" pageId="3" pageNumber="4">Samburupithecus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF990F5FFB71FA568EE7FA59" authorityName="von Königswald" authorityYear="1972" box="[1266,1417,1503,1524]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFB71FA568EE7FA59" box="[1266,1417,1503,1524]" italics="true" pageId="3" pageNumber="4">Graecopithecus</emphasis>
</taxonomicName>
) all recover a clade that includes
<taxonomicName id="4C7D4D72FF990F5FFBE5FA728F8AF9BD" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[1126,1252,1531,1552]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFBE5FA728F8AF9BD" box="[1126,1252,1531,1552]" italics="true" pageId="3" pageNumber="4">Anadoluvius</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF990F5FFAA2FA728ED1F9BD" box="[1313,1471,1531,1552]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFAA2FA728ED1F9BD" box="[1313,1471,1531,1552]" italics="true" pageId="3" pageNumber="4">Ouranopithecus</emphasis>
</taxonomicName>
as sister taxa, which in turn is either the sister taxon to the dryopithecins or in an unresolved polychotomy with the dryopithecins and the crown hominines.
<taxonomicName id="4C7D4D72FF990F5FFB5DF9C68E1CF9C9" authorityName="Kunimatsu, Nakatsukasa, Sawada, Sakai, Hyodo, Hyodo, Itaya, Nakaya, Saegusa, Mazurier, Saneyoshi, Tsujikawa, Yamamoto &amp; Mbua" authorityYear="2007" box="[1246,1394,1615,1636]" class="Mammalia" family="Hominidae" genus="Nakalipithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFB5DF9C68E1CF9C9" box="[1246,1394,1615,1636]" italics="true" pageId="3" pageNumber="4">Nakalipithecus</emphasis>
</taxonomicName>
, which has been interpreted as a potential ancestor of
<taxonomicName id="4C7D4D72FF990F5FFA87F9E28ECCF92D" box="[1284,1442,1643,1664]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFA87F9E28ECCF92D" box="[1284,1442,1643,1664]" italics="true" pageId="3" pageNumber="4">Ouranopithecus</emphasis>
</taxonomicName>
, is outside the crown hominids in this analysis, as suggested elsewhere
<bibRefCitation id="EFEC4B00FF990F5FFC08F91788FBF91C" author="Begun, D. R." box="[907,917,1694,1713]" editor="Henke, W. &amp; Tattersall, I." journalOrPublisher="Springer" pageId="3" pageNumber="4" pagination="1261 - 1332" refId="ref9660" refString="1. Begun, D. R. Fossil Record of Miocene Hominoids in Handbook of Paleoanthropology (eds. Henke, W. &amp; Tattersall, I.) 1261 - 1332 (Springer, 2015)." title="Fossil Record of Miocene Hominoids" type="book chapter" volumeTitle="Handbook of Paleoanthropology" year="2015">1</bibRefCitation>
,
<bibRefCitation id="EFEC4B00FF990F5FFC1AF91788C2F91C" author="Ward, S. C. &amp; Kimbel, W. H." box="[921,940,1694,1713]" journalOrPublisher="Am. J. Phys. Anthropol." pageId="3" pageNumber="4" pagination="157 - 171" part="61" refId="ref10832" refString="31. Ward, S. C. &amp; Kimbel, W. H. Subnasal alveolar morphology and the systemic postion of Sivapithecus. Am. J. Phys. Anthropol. 61, 157 - 171 (1983)." title="Subnasal alveolar morphology and the systemic postion of Sivapithecus" type="journal article" year="1983">31</bibRefCitation>
<taxonomicName id="4C7D4D72FF990F5FFC35F92B8F3CF91A" box="[950,1106,1698,1719]" class="Mammalia" family="Hominidae" genus="Ankarapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFC35F92B8F3CF91A" box="[950,1106,1698,1719]" italics="true" pageId="3" pageNumber="4">Ankarapithecus</emphasis>
</taxonomicName>
is identified as a pongine and the widely accepted
<taxonomicName id="4C7D4D72FF990F5FFC50F9368895F979" authorityName="Oken" authorityYear="1816" box="[979,1019,1727,1748]" class="Mammalia" family="Hominidae" genus="Pan" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFC50F9368895F979" box="[979,1019,1727,1748]" italics="true" pageId="3" pageNumber="4">Pan</emphasis>
</taxonomicName>
-hominin clade is supported as well.
</paragraph>
<paragraph id="8BC236F1FF990F58FCC0F9538AC1FA74" blockId="3.[810,1472,135,1972]" lastBlockId="4.[116,779,1057,1972]" lastPageId="4" lastPageNumber="5" pageId="3" pageNumber="4">
<emphasis id="B909EAE3FF990F5FFCC0F9538FEBF942" box="[835,1157,1754,1775]" italics="true" pageId="3" pageNumber="4">
<taxonomicName id="4C7D4D72FF990F5FFCC0F953888DF942" box="[835,995,1754,1775]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Ouranopithecus</taxonomicName>
,
<taxonomicName id="4C7D4D72FF990F5FFC6CF9538FEBF942" authorityName="von Königswald" authorityYear="1972" box="[1007,1157,1754,1775]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Graecopithecus</taxonomicName>
</emphasis>
and
<taxonomicName id="4C7D4D72FF990F5FFB38F9538E57F942" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[1211,1337,1754,1775]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFB38F9538E57F942" box="[1211,1337,1754,1775]" italics="true" pageId="3" pageNumber="4">Anadoluvius</emphasis>
</taxonomicName>
share a suite of derived characters of the jaws and dentition that support their status as a distinct clade. Although
<taxonomicName id="4C7D4D72FF990F5FFB1EF89B8E5DF88A" authorityName="von Königswald" authorityYear="1972" box="[1181,1331,1810,1831]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFB1EF89B8E5DF88A" box="[1181,1331,1810,1831]" italics="true" pageId="3" pageNumber="4">Graecopithecus</emphasis>
</taxonomicName>
could not be included in the analyses that yielded well resolved phylogenies, due to its many missing data (90%), all previous analyses of
<taxonomicName id="4C7D4D72FF990F5FFCA9F8EF88AEF8D6" authorityName="von Königswald" authorityYear="1972" box="[810,960,1894,1915]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFCA9F8EF88AEF8D6" box="[810,960,1894,1915]" italics="true" pageId="3" pageNumber="4">Graecopithecus</emphasis>
</taxonomicName>
associate it phylogenetically with
<taxonomicName id="4C7D4D72FF990F5FFCA9F80B88A6F83A" box="[810,968,1922,1943]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF990F5FFCA9F80B88A6F83A" box="[810,968,1922,1943]" italics="true" pageId="3" pageNumber="4">Ouranopithecus</emphasis>
</taxonomicName>
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,
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. The core attributes of the Balkan/ Anatolian late Miocene apes are large, thickly enameled molars, transversely robust mandibles, small canines, and large size. However, among these taxa there is diversity.
<taxonomicName id="4C7D4D72FF9E0F58FDD9FBB4899EFBFF" authorityName="von Königswald" authorityYear="1972" box="[602,752,1085,1106]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FDD9FBB4899EFBFF" box="[602,752,1085,1106]" italics="true" pageId="4" pageNumber="5">Graecopithecus</emphasis>
</taxonomicName>
is distinguished from
<taxonomicName id="4C7D4D72FF9E0F58FEC0FBD08A8FFBC3" box="[323,481,1113,1134]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FEC0FBD08A8FFBC3" box="[323,481,1113,1134]" italics="true" pageId="4" pageNumber="5">Ouranopithecus</emphasis>
</taxonomicName>
(
<emphasis id="B909EAE3FF9E0F58FE76FBD0895BFBC3" box="[501,565,1113,1134]" italics="true" pageId="4" pageNumber="5">contra</emphasis>
35) in its relatively large M
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compared with both the M
<subScript id="17F934B4FF9E0F58FE7CFBF78966FB21" attach="left" box="[511,520,1150,1164]" fontSize="6" pageId="4" pageNumber="5">1</subScript>
and corpus breadth, its more vertical mandibular symphysis and in details of root morphology
<superScript id="7C089BB9FF9E0F58FF73FB2E8A47FB17" attach="left" box="[240,297,1191,1210]" fontSize="8" pageId="4" pageNumber="5">
<bibRefCitation id="EFEC4B00FF9E0F58FF73FB2E8B94FB17" author="Begun, D. R." box="[240,250,1191,1210]" editor="Henke, W. &amp; Tattersall, I." journalOrPublisher="Springer" pageId="4" pageNumber="5" pagination="1261 - 1332" refId="ref9660" refString="1. Begun, D. R. Fossil Record of Miocene Hominoids in Handbook of Paleoanthropology (eds. Henke, W. &amp; Tattersall, I.) 1261 - 1332 (Springer, 2015)." title="Fossil Record of Miocene Hominoids" type="book chapter" volumeTitle="Handbook of Paleoanthropology" year="2015">1</bibRefCitation>
,
<bibRefCitation id="EFEC4B00FF9E0F58FF7DFB2E8A7FFB17" author="Fuss, J. &amp; Spassov, N. &amp; Begun, D. R. &amp; Bohme, M." box="[254,273,1191,1210]" journalOrPublisher="PLoS ONE" pageId="4" pageNumber="5" pagination="0177127" part="12" refId="ref10011" refString="11. Fuss, J., Spassov, N., Begun, D. R. &amp; Bohme, M. Potential hominin affinities of Graecopithecus from the Late Miocene of Europe. PLoS ONE 12, e 0177127 (2017)." title="Potential hominin affinities of Graecopithecus from the Late Miocene of Europe" type="journal article" year="2017">11</bibRefCitation>
,
<bibRefCitation id="EFEC4B00FF9E0F58FE95FB2E8A47FB17" author="Koufos, G. D." box="[278,297,1191,1210]" journalOrPublisher="Ann. Paleontol." pageId="4" pageNumber="5" pagination="227 - 240" part="91" refId="ref10201" refString="15. Koufos, G. D. &amp; Bonis, Lde The Late Miocene hominoids Ouranopithecus and Graecopithecus. Implications about their relationships and taxonomy. Ann. Paleontol. 91, 227 - 240 (2005)." title="Bonis, Lde The Late Miocene hominoids Ouranopithecus and Graecopithecus. Implications about their relationships and taxonomy" type="journal article" year="2005">15</bibRefCitation>
</superScript>
<taxonomicName id="4C7D4D72FF9E0F58FEB7FB258ADCFB6C" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[308,434,1196,1217]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FEB7FB258ADCFB6C" box="[308,434,1196,1217]" italics="true" pageId="4" pageNumber="5">Anadoluvius</emphasis>
</taxonomicName>
has the same lower dental root formula (P
<subScript id="17F934B4FF9E0F58FF69FB5B8B9DFB4D" attach="left" box="[234,243,1234,1248]" fontSize="6" pageId="4" pageNumber="5">3</subScript>
to M
<subScript id="17F934B4FF9E0F58FEB9FB5B8A2DFB4D" attach="left" box="[314,323,1234,1248]" fontSize="6" pageId="4" pageNumber="5">1</subScript>
) as
<taxonomicName id="4C7D4D72FF9E0F58FEFEFB41897DFB70" authorityName="von Königswald" authorityYear="1972" box="[381,531,1224,1245]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FEFEFB41897DFB70" box="[381,531,1224,1245]" italics="true" pageId="4" pageNumber="5">Graecopithecus</emphasis>
</taxonomicName>
and both differ from
<taxonomicName id="4C7D4D72FF9E0F58FFF7FB6D8A7CFB54" box="[116,274,1252,1273]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FFF7FB6D8A7CFB54" box="[116,274,1252,1273]" italics="true" pageId="4" pageNumber="5">Ouranopithecus</emphasis>
</taxonomicName>
. The frontal bone between the superior orbital margin and the anterior temporal line is preserved in XIR-1 (
<taxonomicName id="4C7D4D72FF9E0F58FFFEFA958A75FA9C" box="[125,283,1308,1329]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FFFEFA958A75FA9C" box="[125,283,1308,1329]" italics="true" pageId="4" pageNumber="5">Ouranopithecus</emphasis>
</taxonomicName>
), which is sufficient to show that it was more vertically oriented than in CO-2100/2800.
<taxonomicName id="4C7D4D72FF9E0F58FDA5FAB189CAFAE0" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[550,676,1336,1357]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FDA5FAB189CAFAE0" box="[550,676,1336,1357]" italics="true" pageId="4" pageNumber="5">Anadoluvius</emphasis>
</taxonomicName>
is further distinguished from
<taxonomicName id="4C7D4D72FF9E0F58FED2FADD8A81FAC4" box="[337,495,1364,1385]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FED2FADD8A81FAC4" box="[337,495,1364,1385]" italics="true" pageId="4" pageNumber="5">Ouranopithecus</emphasis>
</taxonomicName>
and other non-hominin hominines in having mesiodistally shorter canines (as was probably the case in
<taxonomicName id="4C7D4D72FF9E0F58FED4FA058A83FA0C" authorityName="von Königswald" authorityYear="1972" box="[343,493,1420,1441]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FED4FA058A83FA0C" box="[343,493,1420,1441]" italics="true" pageId="4" pageNumber="5">Graecopithecus</emphasis>
</taxonomicName>
based on canine root size and shape) that lack mesial grooves and lingual cingula (unknown for
<taxonomicName id="4C7D4D72FF9E0F58FE88FA4A8ACFFA75" authorityName="von Königswald" authorityYear="1972" box="[267,417,1475,1496]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FE88FA4A8ACFFA75" box="[267,417,1475,1496]" italics="true" pageId="4" pageNumber="5">Graecopithecus</emphasis>
</taxonomicName>
).
</paragraph>
<caption id="DF026679FF9E0F58FFF7FD548844FC46" ID-DOI="http://doi.org/10.5281/zenodo.8279665" ID-Zenodo-Dep="8279665" httpUri="https://zenodo.org/record/8279665/files/figure.png" pageId="4" pageNumber="5" startId="4.[116,146,733,752]" targetBox="[156,1428,137,704]" targetPageId="4" targetType="figure">
<paragraph id="8BC236F1FF9E0F58FFF7FD548844FC46" blockId="4.[116,1472,733,1003]" pageId="4" pageNumber="5">
<emphasis id="B909EAE3FF9E0F58FFF7FD548AD9FD5D" bold="true" box="[116,439,733,752]" pageId="4" pageNumber="5">Fig. 4 Strict consensus cladograms.</emphasis>
The four taxon sets each produced cladograms with the same topology whether character states were left unordered or a subset were ordered (see Methods and Supplementary Note 5 for details).
<emphasis id="B909EAE3FF9E0F58FCC6FD70883EFCA1" bold="true" box="[837,848,761,780]" pageId="4" pageNumber="5">a</emphasis>
18 OTUs. The four taxa with the fewest codable character states (
<taxonomicName id="4C7D4D72FF9E0F58FFFEFC9D8B95FC85" authorityName="von Königswald" authorityYear="1972" box="[125,251,788,808]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FFFEFC9D8B95FC85" box="[125,251,788,808]" italics="true" pageId="4" pageNumber="5">Graecopithecus</emphasis>
</taxonomicName>
, 10%,
<taxonomicName id="4C7D4D72FF9E0F58FEB5FC9D8AD5FC85" authorityName="Suwa, Kono, Katoh, Asfaw &amp; Beyene" authorityYear="2007" box="[310,443,788,808]" class="Mammalia" genus="Chororapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FEB5FC9D8AD5FC85" box="[310,443,788,808]" italics="true" pageId="4" pageNumber="5">Chororapithecus</emphasis>
</taxonomicName>
,13%,
<taxonomicName id="4C7D4D72FF9E0F58FE6CFC9D8912FC85" authorityName="Ishida &amp; Pickford" authorityYear="1997" box="[495,636,788,808]" class="Mammalia" genus="Samburupithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FE6CFC9D8912FC85" box="[495,636,788,808]" italics="true" pageId="4" pageNumber="5">Samburupithecus</emphasis>
</taxonomicName>
, 18%, and
<taxonomicName id="4C7D4D72FF9E0F58FD5FFC9D8879FC85" authorityName="Senut, Pickford, Gommery, Mein, Cheboi &amp; Coppens" authorityYear="2001" box="[732,791,788,808]" class="Mammalia" family="Hominidae" genus="Orrorin" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FD5FFC9D8879FC85" box="[732,791,788,808]" italics="true" pageId="4" pageNumber="5">Orrorin</emphasis>
</taxonomicName>
, 29%) were excluded, as was
<taxonomicName id="4C7D4D72FF9E0F58FBA0FC9D8FCCFC85" box="[1059,1186,788,808]" class="Mammalia" family="Hominidae" genus="Sahelanthropus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FBA0FC9D8FCCFC85" box="[1059,1186,788,808]" italics="true" pageId="4" pageNumber="5">Sahelanthropus</emphasis>
</taxonomicName>
. Both
<taxonomicName id="4C7D4D72FF9E0F58FB59FC9D8E7BFC85" authorityName="Senut, Pickford, Gommery, Mein, Cheboi &amp; Coppens" authorityYear="2001" box="[1242,1301,788,808]" class="Mammalia" family="Hominidae" genus="Orrorin" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FB59FC9D8E7BFC85" box="[1242,1301,788,808]" italics="true" pageId="4" pageNumber="5">Orrorin</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF9E0F58FAC2FC9D8EAEFC85" box="[1345,1472,788,808]" class="Mammalia" family="Hominidae" genus="Sahelanthropus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FAC2FC9D8EAEFC85" box="[1345,1472,788,808]" italics="true" pageId="4" pageNumber="5">Sahelanthropus</emphasis>
</taxonomicName>
were coded from published descriptions, which introduces uncertainty (DRB, who coded all characters in this analysis, was unable to code characters from these taxa through direct observation).
<emphasis id="B909EAE3FF9E0F58FE52FCC58AB2FCF2" bold="true" box="[465,476,844,863]" pageId="4" pageNumber="5">b</emphasis>
19 OTUs, with
<taxonomicName id="4C7D4D72FF9E0F58FDE7FCC5898CFCCD" box="[612,738,844,864]" class="Mammalia" family="Hominidae" genus="Sahelanthropus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FDE7FCC5898CFCCD" box="[612,738,844,864]" italics="true" pageId="4" pageNumber="5">Sahelanthropus</emphasis>
</taxonomicName>
added.
<emphasis id="B909EAE3FF9E0F58FCABFCC5885CFCF2" bold="true" box="[808,818,844,863]" pageId="4" pageNumber="5">c</emphasis>
20 OTUs with
<taxonomicName id="4C7D4D72FF9E0F58FC39FCC5889BFCCD" authorityName="Senut, Pickford, Gommery, Mein, Cheboi &amp; Coppens" authorityYear="2001" box="[954,1013,844,864]" class="Mammalia" family="Hominidae" genus="Orrorin" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FC39FCC5889BFCCD" box="[954,1013,844,864]" italics="true" pageId="4" pageNumber="5">Orrorin</emphasis>
</taxonomicName>
. There is a decrease in resolution with the inclusion of
<taxonomicName id="4C7D4D72FF9E0F58FF0EFCE18A65FCD1" box="[141,267,872,892]" class="Mammalia" family="Hominidae" genus="Sahelanthropus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FF0EFCE18A65FCD1" box="[141,267,872,892]" italics="true" pageId="4" pageNumber="5">Sahelanthropus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF9E0F58FEBAFCE18A1AFCD1" authorityName="Senut, Pickford, Gommery, Mein, Cheboi &amp; Coppens" authorityYear="2001" box="[313,372,872,892]" class="Mammalia" family="Hominidae" genus="Orrorin" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FEBAFCE18A1AFCD1" box="[313,372,872,892]" italics="true" pageId="4" pageNumber="5">Orrorin</emphasis>
</taxonomicName>
but the tree topologies are otherwise consistent.
<emphasis id="B909EAE3FF9E0F58FCB6FCE188D4FCD1" box="[821,954,872,892]" italics="true" pageId="4" pageNumber="5">Sahelantthropus</emphasis>
is always recovered as a stem hominid and
<taxonomicName id="4C7D4D72FF9E0F58FACFFCE18EE9FCD1" authorityName="Senut, Pickford, Gommery, Mein, Cheboi &amp; Coppens" authorityYear="2001" box="[1356,1415,872,892]" class="Mammalia" family="Hominidae" genus="Orrorin" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FACFFCE18EE9FCD1" box="[1356,1415,872,892]" italics="true" pageId="4" pageNumber="5">Orrorin</emphasis>
</taxonomicName>
as a hominin. The first three cladograms all recover a hominine clade that includes the thickly enameled Balkan taxa and the dryopithecins.
<emphasis id="B909EAE3FF9E0F58FA84FC0D8E7CFC3A" bold="true" box="[1287,1298,900,919]" pageId="4" pageNumber="5">d</emphasis>
23 OTUs, including all taxa. Little resolution remains among hominids, with recognized clades (pongines) unresolved. This cladogram also fails to recover
<taxonomicName id="4C7D4D72FF9E0F58FA96FC298EF8FC19" box="[1301,1430,928,948]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FA96FC298EF8FC19" box="[1301,1430,928,948]" italics="true" pageId="4" pageNumber="5">Ouranopithecus</emphasis>
</taxonomicName>
as a hominine, which is otherwise a common result in previous analyses. Bremer support values, character states, character definitions and the character matrix (nexus) are all included in Supplementary Note 5 and Supplementary Data 3.
</paragraph>
</caption>
<paragraph id="8BC236F1FF9E0F58FF0DFA698FB9FA9F" blockId="4.[116,779,1057,1972]" lastBlockId="4.[810,1472,1057,1972]" pageId="4" pageNumber="5">
In quantitative attributes, the PCA (Supplementary Fig. 6) illustrates the overall distinctiveness of
<taxonomicName id="4C7D4D72FF9E0F58FD98FA7289F7F9BD" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[539,665,1531,1552]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FD98FA7289F7F9BD" box="[539,665,1531,1552]" italics="true" pageId="4" pageNumber="5">Anadoluvius</emphasis>
</taxonomicName>
compared with Balkan apes. The male
<taxonomicName id="4C7D4D72FF9E0F58FE0DF99E8962F981" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[398,524,1559,1580]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FE0DF99E8962F981" box="[398,524,1559,1580]" italics="true" pageId="4" pageNumber="5">Anadoluvius</emphasis>
</taxonomicName>
mandible CO-300/305 is distinguished from male
<taxonomicName id="4C7D4D72FF9E0F58FE1EF9BA8955F9E5" box="[413,571,1587,1608]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FE1EF9BA8955F9E5" box="[413,571,1587,1608]" italics="true" pageId="4" pageNumber="5">Ouranopithecus</emphasis>
</taxonomicName>
in many metric comparisons (canine-M
<bibRefCitation id="EFEC4B00FF9E0F58FEE9F9D08A1DF9CA" author="Begun, D. R. &amp; Nargolwalla, M. C. &amp; Kordos, L." box="[362,371,1625,1639]" journalOrPublisher="Evol. Anthropol." pageId="4" pageNumber="5" pagination="10 - 23" part="21" refId="ref9699" refString="2. Begun, D. R., Nargolwalla, M. C. &amp; Kordos, L. European Miocene hominids and the origin of the African ape and human clade. Evol. Anthropol. 21, 10 - 23 (2002)." title="European Miocene hominids and the origin of the African ape and human clade" type="journal article" year="2002">
<subScript id="17F934B4FF9E0F58FEE9F9D08A1DF9CA" attach="left" box="[362,371,1625,1639]" fontSize="6" pageId="4" pageNumber="5">2</subScript>
</bibRefCitation>
ratio, canine/geometric mean, relative mandibular breadth, symphyseal-molar distance, relative P
<subScript id="17F934B4FF9E0F58FC82F9FD8864F92F" attach="left" box="[769,778,1652,1666]" fontSize="6" pageId="4" pageNumber="5">4</subScript>
length and size, and relative P
<subScript id="17F934B4FF9E0F58FE2AF9198ADCF933" attach="left" box="[425,434,1680,1694]" fontSize="6" pageId="4" pageNumber="5">4</subScript>
, M
<subScript id="17F934B4FF9E0F58FE55F9198AB1F933" attach="left" box="[470,479,1680,1694]" fontSize="6" pageId="4" pageNumber="5">1</subScript>
and M
<subScript id="17F934B4FF9E0F58FDA8F919895AF933" attach="left" box="[555,564,1680,1694]" fontSize="6" pageId="4" pageNumber="5">2</subScript>
size (Supplementary Fig. 8af).
<taxonomicName id="4C7D4D72FF9E0F58FF60F92B8A0FF91A" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[227,353,1698,1719]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FF60F92B8A0FF91A" box="[227,353,1698,1719]" italics="true" pageId="4" pageNumber="5">Anadoluvius</emphasis>
</taxonomicName>
is distinguished from male
<taxonomicName id="4C7D4D72FF9E0F58FD01F92B8BFDF979" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FD01F92B8BFDF979" italics="true" pageId="4" pageNumber="5">Ouranopithecus</emphasis>
</taxonomicName>
in canine size relative to postcanine tooth size at every tooth position for both upper and lower tooth rows (Supplementary Fig. 11).
<taxonomicName id="4C7D4D72FF9E0F58FF5AF97F8A39F8A6" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[217,343,1782,1803]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FF5AF97F8A39F8A6" box="[217,343,1782,1803]" italics="true" pageId="4" pageNumber="5">Anadoluvius</emphasis>
</taxonomicName>
is distinguished from
<taxonomicName id="4C7D4D72FF9E0F58FDD1F97F8986F8A6" authorityName="von Königswald" authorityYear="1972" box="[594,744,1782,1803]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FDD1F97F8986F8A6" box="[594,744,1782,1803]" italics="true" pageId="4" pageNumber="5">Graecopithecus</emphasis>
</taxonomicName>
in relative mandibular breadth, symphyseal-molar distance, P
<subScript id="17F934B4FF9E0F58FC82F8958864F887" attach="left" box="[769,778,1820,1834]" fontSize="6" pageId="4" pageNumber="5">4</subScript>
length, M
<subScript id="17F934B4FF9E0F58FF59F8B18B8DF8EB" attach="left" box="[218,227,1848,1862]" fontSize="6" pageId="4" pageNumber="5">2</subScript>
relative size and M
<subScript id="17F934B4FF9E0F58FE3BF8B18AAFF8EB" attach="left" box="[440,449,1848,1862]" fontSize="6" pageId="4" pageNumber="5">2</subScript>
size relative to corpus breadth (Supplementary Fig. 8). In addition, the mandibular arch is wider and relative mandibular breadth larger at every dental position in
<taxonomicName id="4C7D4D72FF9E0F58FFF7F80B8B9CF83A" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[116,242,1922,1943]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FFF7F80B8B9CF83A" box="[116,242,1922,1943]" italics="true" pageId="4" pageNumber="5">Anadoluvius</emphasis>
</taxonomicName>
(Supplementary Fig. 7). Unfortunately,
<taxonomicName id="4C7D4D72FF9E0F58FD0EF80B8BFDF81E" authorityName="Kunimatsu, Nakatsukasa, Sawada, Sakai, Hyodo, Hyodo, Itaya, Nakaya, Saegusa, Mazurier, Saneyoshi, Tsujikawa, Yamamoto &amp; Mbua" authorityYear="2007" class="Mammalia" family="Hominidae" genus="Nakalipithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FD0EF80B8BFDF81E" italics="true" pageId="4" pageNumber="5">Nakalipithecus</emphasis>
</taxonomicName>
is insufficiently preserved to be included in most of these quantitative analyses, which are scaled using the geometric mean.
<taxonomicName id="4C7D4D72FF9E0F58FCA9FBB488D0FBFF" authorityName="Kunimatsu, Nakatsukasa, Sawada, Sakai, Hyodo, Hyodo, Itaya, Nakaya, Saegusa, Mazurier, Saneyoshi, Tsujikawa, Yamamoto &amp; Mbua" authorityYear="2007" box="[810,958,1085,1106]" class="Mammalia" family="Hominidae" genus="Nakalipithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FCA9FBB488D0FBFF" box="[810,958,1085,1106]" italics="true" pageId="4" pageNumber="5">Nakalipithecus</emphasis>
</taxonomicName>
lacks one or more of the quantitative attributes needed to generate the geometric mean used in this analysis. We ran an analysis of M
<subScript id="17F934B4FF9E0F58FC7EFBF78F68FB21" attach="left" box="[1021,1030,1150,1164]" fontSize="6" pageId="4" pageNumber="5">2</subScript>
size relative to mandibular corpus breadth at the level of mid M
<subScript id="17F934B4FF9E0F58FB85FB138F61FB05" attach="left" box="[1030,1039,1178,1192]" fontSize="6" pageId="4" pageNumber="5">1</subScript>
, which are data that have been published for
<taxonomicName id="4C7D4D72FF9E0F58FCD7FB258886FB6C" authorityName="Kunimatsu, Nakatsukasa, Sawada, Sakai, Hyodo, Hyodo, Itaya, Nakaya, Saegusa, Mazurier, Saneyoshi, Tsujikawa, Yamamoto &amp; Mbua" authorityYear="2007" box="[852,1000,1196,1217]" class="Mammalia" family="Hominidae" genus="Nakalipithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FCD7FB258886FB6C" box="[852,1000,1196,1217]" italics="true" pageId="4" pageNumber="5">Nakalipithecus</emphasis>
</taxonomicName>
. In this ratio the
<taxonomicName id="4C7D4D72FF9E0F58FB39FB258E20FB6C" authorityName="Kunimatsu, Nakatsukasa, Sawada, Sakai, Hyodo, Hyodo, Itaya, Nakaya, Saegusa, Mazurier, Saneyoshi, Tsujikawa, Yamamoto &amp; Mbua" authorityYear="2007" box="[1210,1358,1196,1217]" class="Mammalia" family="Hominidae" genus="Nakalipithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FB39FB258E20FB6C" box="[1210,1358,1196,1217]" italics="true" pageId="4" pageNumber="5">Nakalipithecus</emphasis>
</taxonomicName>
specimen, which is probably male, falls within the range of gorilla females,
<taxonomicName id="4C7D4D72FF9E0F58FCA9FB6C883CFB57" authorityName="Oken" authorityYear="1816" box="[810,850,1253,1274]" class="Mammalia" family="Hominidae" genus="Pan" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FCA9FB6C883CFB57" box="[810,850,1253,1274]" italics="true" pageId="4" pageNumber="5">Pan</emphasis>
</taxonomicName>
of both sexes, and female
<taxonomicName id="4C7D4D72FF9E0F58FBE0FB6D8E6FFB54" box="[1123,1281,1252,1273]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FBE0FB6D8E6FFB54" box="[1123,1281,1252,1273]" italics="true" pageId="4" pageNumber="5">Ouranopithecus</emphasis>
</taxonomicName>
, outside the range of gorilla males,
<taxonomicName id="4C7D4D72FF9E0F58FC62FA898F11FAB8" box="[993,1151,1280,1301]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FC62FA898F11FAB8" box="[993,1151,1280,1301]" italics="true" pageId="4" pageNumber="5">Ouranopithecus</emphasis>
</taxonomicName>
males, Nikiti,
<taxonomicName id="4C7D4D72FF9E0F58FAA0FA898ED7FAB8" authorityName="von Königswald" authorityYear="1972" box="[1315,1465,1280,1301]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FAA0FA898ED7FAB8" box="[1315,1465,1280,1301]" italics="true" pageId="4" pageNumber="5">Graecopithecus</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C7D4D72FF9E0F58FCDBFA9588B8FA9C" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[856,982,1308,1329]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FCDBFA9588B8FA9C" box="[856,982,1308,1329]" italics="true" pageId="4" pageNumber="5">Anadoluvius</emphasis>
</taxonomicName>
(Supplementary Fig. 8f).
</paragraph>
<paragraph id="8BC236F1FF9E0F58FCC0FAB18F81FA74" blockId="4.[810,1472,1057,1972]" pageId="4" pageNumber="5">
<taxonomicName id="4C7D4D72FF9E0F58FCC0FAB1888FFAE0" box="[835,993,1336,1357]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FCC0FAB1888FFAE0" box="[835,993,1336,1357]" italics="true" pageId="4" pageNumber="5">Ouranopithecus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF9E0F58FBA1FAB18FCEFAE0" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[1058,1184,1336,1357]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FBA1FAB18FCEFAE0" box="[1058,1184,1336,1357]" italics="true" pageId="4" pageNumber="5">Anadoluvius</emphasis>
</taxonomicName>
lack shared derived characters of the pongines (greatly elongated premaxilla substantially or completely overlapping the maxillary palatine process, expanded zygoma, tall orbits, narrow interorbital space, reduced or absent ethmoidal frontal sinus, circumorbital costae.) There is no evidence for their inclusion in Ponginae.
</paragraph>
<paragraph id="8BC236F1FF9E0F58FCC0FA568F79F8E9" blockId="4.[810,1472,1057,1972]" pageId="4" pageNumber="5">
<taxonomicName id="4C7D4D72FF9E0F58FCC0FA56888FFA59" box="[835,993,1503,1524]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FCC0FA56888FFA59" box="[835,993,1503,1524]" italics="true" pageId="4" pageNumber="5">Ouranopithecus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF9E0F58FBA4FA568FCBFA59" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[1063,1189,1503,1524]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FBA4FA568FCBFA59" box="[1063,1189,1503,1524]" italics="true" pageId="4" pageNumber="5">Anadoluvius</emphasis>
</taxonomicName>
share with dryopithecins (European middle and late Miocene apes with affinities to
<taxonomicName id="4C7D4D72FF9E0F58FCA9F99E88C2F981" authorityName="Lartet" authorityYear="1856" box="[810,940,1559,1580]" class="Mammalia" family="Hominidae" genus="Dryopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FCA9F99E88C2F981" box="[810,940,1559,1580]" italics="true" pageId="4" pageNumber="5">Dryopithecus</emphasis>
</taxonomicName>
) a series of characters found among hominines
<bibRefCitation id="EFEC4B00FF9E0F58FA22F99B8EC5F988" author="Begun, D. R." box="[1441,1451,1554,1573]" editor="Henke, W. &amp; Tattersall, I." journalOrPublisher="Springer" pageId="4" pageNumber="5" pagination="1261 - 1332" refId="ref9660" refString="1. Begun, D. R. Fossil Record of Miocene Hominoids in Handbook of Paleoanthropology (eds. Henke, W. &amp; Tattersall, I.) 1261 - 1332 (Springer, 2015)." title="Fossil Record of Miocene Hominoids" type="book chapter" volumeTitle="Handbook of Paleoanthropology" year="2015">1</bibRefCitation>
<bibRefCitation id="EFEC4B00FF9E0F58FA36F99B8ED1F988" author="Begun, D. R." box="[1461,1471,1554,1573]" journalOrPublisher="Geodiversitas" pageId="4" pageNumber="5" pagination="789 - 816" part="31" refId="ref9746" refString="3. Begun, D. R. Dryopithecus, Darwin, de Bonis and the European origin of the African apes and human clade. Geodiversitas 31, 789 - 816 (2009)." title="Dryopithecus, Darwin, de Bonis and the European origin of the African apes and human clade" type="journal article" year="2009">3</bibRefCitation>
These include a ventrally rotated palate, a stepped subnasal fossa, broad, flat nasal aperture base, short nasal bones, nasal aperture apex superior to the infraorbital margins, robust lateral orbital pillars, frontal sinus expanded below nasion, incipient supraorbital torus, more horizontal frontal squama
<bibRefCitation id="EFEC4B00FF9E0F58FB60F9178F83F91C" author="Begun, D. R." box="[1251,1261,1694,1713]" editor="Henke, W. &amp; Tattersall, I." journalOrPublisher="Springer" pageId="4" pageNumber="5" pagination="1261 - 1332" refId="ref9660" refString="1. Begun, D. R. Fossil Record of Miocene Hominoids in Handbook of Paleoanthropology (eds. Henke, W. &amp; Tattersall, I.) 1261 - 1332 (Springer, 2015)." title="Fossil Record of Miocene Hominoids" type="book chapter" volumeTitle="Handbook of Paleoanthropology" year="2015">1</bibRefCitation>
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The phylogenetic significance of some of these shared attributes is disputed, particularly concerning the dryopithecins
<bibRefCitation id="EFEC4B00FF9E0F58FB55F95C8F8EF945" author="Pugh, K. D." box="[1238,1248,1749,1768]" journalOrPublisher="J. Hum. Evol" pageId="4" pageNumber="5" refId="ref9940" refString="9. Pugh, K. D. Phylogenetic analysis of Middle-Late Miocene apes. J. Hum. Evol. https: // doi. org / 10.1016 / j. jhevol. 2021.103140 (2022)." title="Phylogenetic analysis of Middle-Late Miocene apes" type="book" year="2022">9</bibRefCitation>
,
<bibRefCitation id="EFEC4B00FF9E0F58FB67F95C8F99F945" author="Alba, D. M." box="[1252,1271,1749,1768]" journalOrPublisher="Evol. Anthropol." pageId="4" pageNumber="5" pagination="254 - 269" part="21" refId="ref10575" refString="24. Alba, D. M. Fossil apes from the Valles-Penedes basin. Evol. Anthropol. 21, 254 - 269 (2012)." title="Fossil apes from the Valles-Penedes basin" type="journal article" year="2012">24</bibRefCitation>
. However, there is broad agreement that
<taxonomicName id="4C7D4D72FF9E0F58FBA2F97F8FD1F8A6" box="[1057,1215,1782,1803]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FBA2F97F8FD1F8A6" box="[1057,1215,1782,1803]" italics="true" pageId="4" pageNumber="5">Ouranopithecus</emphasis>
</taxonomicName>
shares enough derived characters with hominines to warrant inclusion in that taxon
<bibRefCitation id="EFEC4B00FF9E0F58FCE1F8A08802F891" author="Begun, D. R." box="[866,876,1833,1852]" editor="Henke, W. &amp; Tattersall, I." journalOrPublisher="Springer" pageId="4" pageNumber="5" pagination="1261 - 1332" refId="ref9660" refString="1. Begun, D. R. Fossil Record of Miocene Hominoids in Handbook of Paleoanthropology (eds. Henke, W. &amp; Tattersall, I.) 1261 - 1332 (Springer, 2015)." title="Fossil Record of Miocene Hominoids" type="book chapter" volumeTitle="Handbook of Paleoanthropology" year="2015">1</bibRefCitation>
<bibRefCitation id="EFEC4B00FF9E0F58FCF5F8A088EEF891" author="Begun, D. R." box="[886,896,1833,1852]" journalOrPublisher="Geodiversitas" pageId="4" pageNumber="5" pagination="789 - 816" part="31" refId="ref9746" refString="3. Begun, D. R. Dryopithecus, Darwin, de Bonis and the European origin of the African apes and human clade. Geodiversitas 31, 789 - 816 (2009)." title="Dryopithecus, Darwin, de Bonis and the European origin of the African apes and human clade" type="journal article" year="2009">3</bibRefCitation>
,
<bibRefCitation id="EFEC4B00FF9E0F58FC06F8A088E1F891" author="Dean, D. &amp; Delson, E." box="[901,911,1833,1852]" journalOrPublisher="Nature" pageId="4" pageNumber="5" pagination="676 - 677" part="359" refId="ref9913" refString="8. Dean, D. &amp; Delson, E. Second gorilla or third chimp? Nature 359, 676 - 677 (1992)." title="Second gorilla or third chimp?" type="journal article" year="1992">8</bibRefCitation>
<bibRefCitation id="EFEC4B00FF9E0F58FC1AF8A088C2F891" author="Ioannidou, M." box="[921,940,1833,1852]" journalOrPublisher="Am. J. Phys. Anthropol." pageId="4" pageNumber="5" pagination="295 - 307" part="170" refId="ref10386" refString="20. Ioannidou, M. A new three-dimensional geometric morphometrics analysis of the Ouranopithecus macedoniensis cranium (Late Miocene, Central Macedonia, Greece). Am. J. Phys. Anthropol. 170, 295 - 307 (2019)." title="A new three-dimensional geometric morphometrics analysis of the Ouranopithecus macedoniensis cranium (Late Miocene, Central Macedonia, Greece)" type="journal article" year="2019">20</bibRefCitation>
(
<tableCitation id="C6FF034AFF9E0F58FC38F8A78F67F8EE" box="[955,1033,1838,1860]" captionStart="Table 1" captionStartId="1.[127,184,1631,1653]" captionTargetBox="[127,767,1722,1956]" captionTargetId="graphics-713@1.[116,777,1609,1966]" captionText="Table 1 Hominine and pongine synapomorphies (18 taxa, unordered)." httpUri="http://table.plazi.org/id/DF026679FF9B0F5DFFFCF9D68B97F93F" pageId="4" pageNumber="5" tableUuid="DF026679FF9B0F5DFFFCF9D68B97F93F">Table 1</tableCitation>
).
</paragraph>
<paragraph id="8BC236F1FF9E0F59FCC0F8C389A8F9E4" blockId="4.[810,1472,1057,1972]" lastBlockId="5.[116,779,1280,1971]" lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5">
The phylogenetic results presented here regarding
<taxonomicName id="4C7D4D72FF9E0F58FAEAF8C38815F8D6" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FAEAF8C38815F8D6" italics="true" pageId="4" pageNumber="5">Ouranopithecus</emphasis>
</taxonomicName>
and its sister,
<taxonomicName id="4C7D4D72FF9E0F58FBA1F8EF8FCEF8D6" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[1058,1184,1894,1915]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9E0F58FBA1F8EF8FCEF8D6" box="[1058,1184,1894,1915]" italics="true" pageId="4" pageNumber="5">Anadoluvius</emphasis>
</taxonomicName>
, are consistent with many previous analyses
<bibRefCitation id="EFEC4B00FF9E0F58FC5AF8F4888DF83D" author="Begun, D. R." box="[985,995,1917,1936]" editor="Henke, W. &amp; Tattersall, I." journalOrPublisher="Springer" pageId="4" pageNumber="5" pagination="1261 - 1332" refId="ref9660" refString="1. Begun, D. R. Fossil Record of Miocene Hominoids in Handbook of Paleoanthropology (eds. Henke, W. &amp; Tattersall, I.) 1261 - 1332 (Springer, 2015)." title="Fossil Record of Miocene Hominoids" type="book chapter" volumeTitle="Handbook of Paleoanthropology" year="2015">1</bibRefCitation>
<bibRefCitation id="EFEC4B00FF9E0F58FC6EF8F48899F83D" author="Begun, D. R." box="[1005,1015,1917,1936]" journalOrPublisher="Geodiversitas" pageId="4" pageNumber="5" pagination="789 - 816" part="31" refId="ref9746" refString="3. Begun, D. R. Dryopithecus, Darwin, de Bonis and the European origin of the African apes and human clade. Geodiversitas 31, 789 - 816 (2009)." title="Dryopithecus, Darwin, de Bonis and the European origin of the African apes and human clade" type="journal article" year="2009">3</bibRefCitation>
,
<bibRefCitation id="EFEC4B00FF9E0F58FC78F8F48F6BF83D" author="Dean, D. &amp; Delson, E." box="[1019,1029,1917,1936]" journalOrPublisher="Nature" pageId="4" pageNumber="5" pagination="676 - 677" part="359" refId="ref9913" refString="8. Dean, D. &amp; Delson, E. Second gorilla or third chimp? Nature 359, 676 - 677 (1992)." title="Second gorilla or third chimp?" type="journal article" year="1992">8</bibRefCitation>
<bibRefCitation id="EFEC4B00FF9E0F58FB93F8F48F4DF83D" author="Ioannidou, M." box="[1040,1059,1917,1936]" journalOrPublisher="Am. J. Phys. Anthropol." pageId="4" pageNumber="5" pagination="295 - 307" part="170" refId="ref10386" refString="20. Ioannidou, M. A new three-dimensional geometric morphometrics analysis of the Ouranopithecus macedoniensis cranium (Late Miocene, Central Macedonia, Greece). Am. J. Phys. Anthropol. 170, 295 - 307 (2019)." title="A new three-dimensional geometric morphometrics analysis of the Ouranopithecus macedoniensis cranium (Late Miocene, Central Macedonia, Greece)" type="journal article" year="2019">20</bibRefCitation>
They are robust in terms of the number of synapomorphies and Bremer support values for many clades (Supplementary
<figureCitation id="13462A74FF9F0F59FE9EFA898AFAFABB" box="[285,404,1280,1302]" captionStart="Figs" captionStartId="3.[810,853,136,158]" captionTargetBox="[129,765,138,1317]" captionTargetId="figure-817@3.[127,767,135,1319]" captionTargetPageId="3" captionText="Figs. 1214; Supplementary Data 2, 3; Supplementary Note 5). All four cladograms are strict consensus. The analysis was run both with all characters unordered and 21 of the 112 characters ordered (see Methods and Supplementary Note 5 for character matrix assumptions). Both analyses yield similar results. The tree topologies are identical, and predictably the tree values are lower in the analyses using ordered characters. All but the cladograms that include the taxa with more than 80% missing data recover a clade that includes Eurasian apes and hominines to the exclusion of pongines and stem hominids (Fig. 4). The potential problem of using data from published sources as opposed to direct observation is illustrated in the results for Sahelanthropus. This taxon, universally attributed to the Homininae and most commonly to the Hominini, is never recovered in these positions in these analyses, being consistently a stem hominid. Sahelanthropus could be coded for 72% of the characters. The potential for coding inconsistent with the criteria used to code other taxa is therefore larger than for Orrorin, which is also coded from the literature, but for which only 29% of the characters could be coded. We mapped synapomorphies and Bremer support values onto two consensus cladograms (Supplementary Figs. 12, 13). Table 1 lists the hominine synapomorphies of the cladogram (ordered and unordered) with the fewest missing data (18 OTUs). A phylogeny consistent with a large majority of the cladograms presented here appears in Fig. 5. Supplementary Fig. 14 shows the results of the analyses of all four taxon sets with all character states ordered. Unlike the unordered and partly ordered analyses, the cladograms with differing OTUs vary widely. The 19 OTU (including Sahelanthropus) fully ordered analysis is consistent with previous ones while the 18 and 20 OTU analyses result in a pongine clade including all European and Anatolian taxa, which contrasts with all previous cladistic results from analyses with large data sets and numerical cladistic methods (e.g. TNT, PAUP, etc). The 20 OTU analysis includes several other unconventional results such as an Orrorin-Anadoluvius clade that, along with Ouranopithecus as the sister to pongines, and a pongine clade that includes all European and Anatolian taxa. The 23 OTU analysis remains highly unresolved. Given the diversity of results of these all ordered analyses and the broader consistency of the analyses in which the data matrices were either unordered or partly ordered, we consider the latter to be more reliable (see Methods for a discussion of our rationale on ordering character states in this analysis)." pageId="5" pageNumber="6">Figs. 12, 13</figureCitation>
). The results also recover the widely accepted relations among crown hominids and hominines and relations within all fossil clades. The Bremer support values for the hominid clade as defined here (15-16) are extremely strong. The hominine clade as defined here is moderate to strong (2-3), and very strong (4 to 8) for the pongine clade (Supplementary
<figureCitation id="13462A74FF9F0F59FFF7FA2E8B96FA10" box="[116,248,1447,1469]" captionStart="Figs" captionStartId="3.[810,853,136,158]" captionTargetBox="[129,765,138,1317]" captionTargetId="figure-817@3.[127,767,135,1319]" captionTargetPageId="3" captionText="Figs. 1214; Supplementary Data 2, 3; Supplementary Note 5). All four cladograms are strict consensus. The analysis was run both with all characters unordered and 21 of the 112 characters ordered (see Methods and Supplementary Note 5 for character matrix assumptions). Both analyses yield similar results. The tree topologies are identical, and predictably the tree values are lower in the analyses using ordered characters. All but the cladograms that include the taxa with more than 80% missing data recover a clade that includes Eurasian apes and hominines to the exclusion of pongines and stem hominids (Fig. 4). The potential problem of using data from published sources as opposed to direct observation is illustrated in the results for Sahelanthropus. This taxon, universally attributed to the Homininae and most commonly to the Hominini, is never recovered in these positions in these analyses, being consistently a stem hominid. Sahelanthropus could be coded for 72% of the characters. The potential for coding inconsistent with the criteria used to code other taxa is therefore larger than for Orrorin, which is also coded from the literature, but for which only 29% of the characters could be coded. We mapped synapomorphies and Bremer support values onto two consensus cladograms (Supplementary Figs. 12, 13). Table 1 lists the hominine synapomorphies of the cladogram (ordered and unordered) with the fewest missing data (18 OTUs). A phylogeny consistent with a large majority of the cladograms presented here appears in Fig. 5. Supplementary Fig. 14 shows the results of the analyses of all four taxon sets with all character states ordered. Unlike the unordered and partly ordered analyses, the cladograms with differing OTUs vary widely. The 19 OTU (including Sahelanthropus) fully ordered analysis is consistent with previous ones while the 18 and 20 OTU analyses result in a pongine clade including all European and Anatolian taxa, which contrasts with all previous cladistic results from analyses with large data sets and numerical cladistic methods (e.g. TNT, PAUP, etc). The 20 OTU analysis includes several other unconventional results such as an Orrorin-Anadoluvius clade that, along with Ouranopithecus as the sister to pongines, and a pongine clade that includes all European and Anatolian taxa. The 23 OTU analysis remains highly unresolved. Given the diversity of results of these all ordered analyses and the broader consistency of the analyses in which the data matrices were either unordered or partly ordered, we consider the latter to be more reliable (see Methods for a discussion of our rationale on ordering character states in this analysis)." pageId="5" pageNumber="6">Figs. 12, 13</figureCitation>
). European and eastern Mediterranean apes are classified as hominine in three of four consensus cladograms. It fails to be resolved only in the analysis of all 23 taxa, including
<emphasis id="B909EAE3FF9F0F59FFF7FA728AB8F9BD" box="[116,470,1531,1552]" italics="true" pageId="5" pageNumber="6">
<taxonomicName id="4C7D4D72FF9F0F59FFF7FA728A4CF9BD" authorityName="Ishida &amp; Pickford" authorityYear="1997" box="[116,290,1531,1552]" class="Mammalia" genus="Samburupithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Samburupithecus</taxonomicName>
,
<taxonomicName id="4C7D4D72FF9F0F59FEB1FA728AB8F9BD" authorityName="Suwa, Kono, Katoh, Asfaw &amp; Beyene" authorityYear="2007" box="[306,470,1531,1552]" class="Mammalia" genus="Chororapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Chororapithecus</taxonomicName>
</emphasis>
and
<taxonomicName id="4C7D4D72FF9F0F59FD97FA7289C4F9BD" authorityName="von Königswald" authorityYear="1972" box="[532,682,1531,1552]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FD97FA7289C4F9BD" box="[532,682,1531,1552]" italics="true" pageId="5" pageNumber="6">Graecopithecus</emphasis>
</taxonomicName>
, missing 82%, 87% and 90% of the data respectively. The latter cladogram also fails to distinguish between pongines and hominines.
</paragraph>
<caption id="DF026679FF9F0F59FFF7FD77883FFB7D" ID-DOI="http://doi.org/10.5281/zenodo.8279667" ID-Zenodo-Dep="8279667" httpUri="https://zenodo.org/record/8279667/files/figure.png" pageId="5" pageNumber="6" startId="5.[116,146,766,785]" targetBox="[236,1352,137,739]" targetPageId="5" targetType="figure">
<paragraph id="8BC236F1FF9F0F59FFF7FD77883FFB7D" blockId="5.[116,1473,766,1232]" pageId="5" pageNumber="6">
<emphasis id="B909EAE3FF9F0F59FFF7FD778F10FCBC" bold="true" box="[116,1150,766,785]" pageId="5" pageNumber="6">Fig. 5 A phylogeny of the taxa included in this analysis consistent with most of the cladograms presented here.</emphasis>
Taxa are positioned in chronological order without regard to geography,with most taxa only known from a limited time span. Exceptions are
<taxonomicName id="4C7D4D72FF9F0F59FC66FC938F4AFC83" authorityName="McNulty, Begun, Kelley, Manthi &amp; Mbua" authorityYear="2015" box="[997,1060,794,814]" class="Mammalia" genus="Ekembo" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FC66FC938F4AFC83" box="[997,1060,794,814]" italics="true" pageId="5" pageNumber="6">Ekembo</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF9F0F59FBCEFC938FDFFC83" box="[1101,1201,794,814]" class="Mammalia" family="Hominidae" genus="Sivapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FBCEFC938FDFFC83" box="[1101,1201,794,814]" italics="true" pageId="5" pageNumber="6">Sivapithecus</emphasis>
</taxonomicName>
, with longer time ranges,which are positioned roughly when they are most abundant, in both cases about mid-way in their known time ranges. The different colored
<emphasis id="B909EAE3FF9F0F59FA87FCBF8E62FCE4" box="[1284,1292,822,841]" italics="true" pageId="5" pageNumber="6"></emphasis>
puddles
<emphasis id="B909EAE3FF9F0F59FAD3FCBF8E36FCE4" box="[1360,1368,822,841]" italics="true" pageId="5" pageNumber="6"></emphasis>
represent hominid clades and/or stages of evolution. These can also be imagined as pools of related species in somewhat delimited space and time with broad ancestordescendant relationships. The lines are disconnected to reflect the difficulty in identifying actual ancestor-descendant relationships, but that these relations can be estimated between
<emphasis id="B909EAE3FF9F0F59FEBFFC038A2AFC30" box="[316,324,906,925]" italics="true" pageId="5" pageNumber="6"></emphasis>
puddles
<emphasis id="B909EAE3FF9F0F59FE04FC038AE1FC30" box="[391,399,906,925]" italics="true" pageId="5" pageNumber="6"></emphasis>
. Blue puddle taxa are stem hominids and are all confined to Africa. Among these taxa the relations of
<taxonomicName id="4C7D4D72FF9F0F59FA8DFC008EF7FC30" authorityName="Ishida &amp; Pickford" authorityYear="1997" box="[1294,1433,905,925]" class="Mammalia" genus="Samburupithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FA8DFC008EF7FC30" box="[1294,1433,905,925]" italics="true" pageId="5" pageNumber="6">Samburupithecus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF9F0F59FFF7FC2C8B99FC14" authorityName="Suwa, Kono, Katoh, Asfaw &amp; Beyene" authorityYear="2007" box="[116,247,933,953]" class="Mammalia" genus="Chororapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FFF7FC2C8B99FC14" box="[116,247,933,953]" italics="true" pageId="5" pageNumber="6">Chororapithecus</emphasis>
</taxonomicName>
are unresolved in the cladograms except in so far as they are excluded from the clade that includes all Eurasian taxa and crown hominins. Other lines of evidence suggest that these taxa are members of the early or middle Miocene radiation of early apes (see text). The orange puddles are the pongines, which probably have their origin within the middle Miocene puddle, although not necessarily any of the taxa included here (another taxon,
<taxonomicName id="4C7D4D72FF9F0F59FAB6FC548EC3FC5C" authorityName="Abel" authorityYear="1903" box="[1333,1453,989,1009]" class="Mammalia" family="Hominidae" genus="Griphopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FAB6FC548EC3FC5C" box="[1333,1453,989,1009]" italics="true" pageId="5" pageNumber="6">Griphopithecus</emphasis>
</taxonomicName>
, known from Europe and Türkiye, would be a member of the blue puddle but was not included in the cladistic analyses).While both are pongines,
<taxonomicName id="4C7D4D72FF9F0F59FAC1FC708ED1FBA0" box="[1346,1471,1017,1037]" class="Mammalia" family="Hominidae" genus="Ankarapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FAC1FC708ED1FBA0" box="[1346,1471,1017,1037]" italics="true" pageId="5" pageNumber="6">Ankarapithecus</emphasis>
</taxonomicName>
lacks derived features shared by
<taxonomicName id="4C7D4D72FF9F0F59FE10FB9C8A96FB84" box="[403,504,1045,1065]" class="Mammalia" family="Hominidae" genus="Sivapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FE10FB9C8A96FB84" box="[403,504,1045,1065]" italics="true" pageId="5" pageNumber="6">Sivapithecus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF9F0F59FDA0FB9C893BFB84" authorityName="Lacepede" authorityYear="1799" box="[547,597,1045,1065]" class="Mammalia" family="Hominidae" genus="Pongo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FDA0FB9C893BFB84" box="[547,597,1045,1065]" italics="true" pageId="5" pageNumber="6">Pongo</emphasis>
</taxonomicName>
, so the line representing the relationship between the latter two bi-passes
<taxonomicName id="4C7D4D72FF9F0F59FB60FB9C8E0EFB84" box="[1251,1376,1045,1065]" class="Mammalia" family="Hominidae" genus="Ankarapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FB60FB9C8E0EFB84" box="[1251,1376,1045,1065]" italics="true" pageId="5" pageNumber="6">Ankarapithecus</emphasis>
</taxonomicName>
. The three green-shade puddles represent the hominine clades as defined here.Bright green are the dryopithecins,with the younger taxa
<taxonomicName id="4C7D4D72FF9F0F59FB24FBB88E7CFBE8" authorityName="Kretzoi" authorityYear="1969" box="[1191,1298,1073,1093]" class="Mammalia" family="Hominidae" genus="Rudapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FB24FBB88E7CFBE8" box="[1191,1298,1073,1093]" italics="true" pageId="5" pageNumber="6">Rudapithecus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF9F0F59FAB8FBB88ED1FBE8" authorityName="Villalta &amp; Crusafont Pairó" authorityYear="1944" box="[1339,1471,1073,1093]" class="Mammalia" family="Hominidae" genus="Hispanopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FAB8FBB88ED1FBE8" box="[1339,1471,1073,1093]" italics="true" pageId="5" pageNumber="6">Hispanopithecus</emphasis>
</taxonomicName>
depicted as closely related and descendant from any of the older taxa or an unknown taxon sharing attributes with these three. The light green puddle includes the Balkan and Anatolian taxa, likely to have descended from somewhere in the dryopithecin puddle.Possible ancestor-descendant relationships are depicted in this puddle. The darker green puddle represents the crown hominines. The various lineages diverge from unknown ancestors, but probably a member of either of the older green shade puddles.Gorillas diverge first, followed by chimpanzees and humans.
<taxonomicName id="4C7D4D72FF9F0F59FC1BFB2988BCFB19" authorityName="Senut, Pickford, Gommery, Mein, Cheboi &amp; Coppens" authorityYear="2001" box="[920,978,1184,1204]" class="Mammalia" family="Hominidae" genus="Orrorin" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FC1BFB2988BCFB19" box="[920,978,1184,1204]" italics="true" pageId="5" pageNumber="6">Orrorin</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF9F0F59FC78FB298F0EFB19" box="[1019,1120,1184,1204]" class="Mammalia" family="Hominidae" genus="Ardipithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FC78FB298F0EFB19" box="[1019,1120,1184,1204]" italics="true" pageId="5" pageNumber="6">Ardipithecus</emphasis>
</taxonomicName>
are depicted in a manner consistent with their sister clade status, without implying a direct ancestor-descendant relationship.
</paragraph>
</caption>
<paragraph id="8BC236F1FF9F0F59FF0DF9C7893DF8D6" blockId="5.[116,779,1280,1971]" pageId="5" pageNumber="6">
The relations among the dryopithecins are consistent with most detailed analyses focused on this group, as are the relations within the Asian clade (
<emphasis id="B909EAE3FF9F0F59FEE9F90F89E0F936" box="[362,654,1670,1691]" italics="true" pageId="5" pageNumber="6">
<taxonomicName id="4C7D4D72FF9F0F59FEE9F90F8966F936" box="[362,520,1670,1691]" class="Mammalia" family="Hominidae" genus="Ankarapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Ankarapithecus</taxonomicName>
,
<taxonomicName id="4C7D4D72FF9F0F59FD97F90F89E0F936" box="[532,654,1670,1691]" class="Mammalia" family="Hominidae" genus="Sivapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Sivapithecus</taxonomicName>
</emphasis>
and
<taxonomicName id="4C7D4D72FF9F0F59FD47F90E886FF931" authorityName="Lacepede" authorityYear="1799" box="[708,769,1671,1692]" class="Mammalia" family="Hominidae" genus="Pongo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FD47F90E886FF931" box="[708,769,1671,1692]" italics="true" pageId="5" pageNumber="6">Pongo</emphasis>
</taxonomicName>
) and the crown hominines
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,
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The consistencies of these cladograms in many details with previous research lend credibility to these results. Recent analyses by (9 and 10) are broadly similar in their results. Among the most parsimonious cladograms reported in ref.
<bibRefCitation id="EFEC4B00FF9F0F59FE33F8848AADF88D" author="Nengo, I. &amp; et al." box="[432,451,1805,1824]" journalOrPublisher="Nature" pageId="5" pageNumber="6" pagination="169 - 174" part="548" refId="ref9979" refString="10. Nengo, I. et al. New infant cranium from the African Miocene sheds light on ape evolution. Nature 548, 169 - 174 (2017)." title="New infant cranium from the African Miocene sheds light on ape evolution" type="journal article" year="2017">10</bibRefCitation>
at least one also recovers a hominine clade that includes European fossil taxa9. also recovers
<taxonomicName id="4C7D4D72FF9F0F59FFF7F8C38A7CF8F2" box="[116,274,1866,1887]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FFF7F8C38A7CF8F2" box="[116,274,1866,1887]" italics="true" pageId="5" pageNumber="6">Ouranopithecus</emphasis>
</taxonomicName>
as a hominine though in that analysis the dryopithecins are recovered as stem hominids.
</paragraph>
<paragraph id="8BC236F1FF9F0F59FF0DF80888ADF8CD" blockId="5.[116,779,1280,1971]" lastBlockId="5.[810,1472,1280,1888]" pageId="5" pageNumber="6">
Other taxa have been linked with
<taxonomicName id="4C7D4D72FF9F0F59FDBCF80889B3F83B" box="[575,733,1921,1942]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FDBCF80889B3F83B" box="[575,733,1921,1942]" italics="true" pageId="5" pageNumber="6">Ouranopithecus</emphasis>
</taxonomicName>
or hominines. Three fossil apes from Africa,
<taxonomicName id="4C7D4D72FF9F0F59FDF2F814886BF81F" authorityName="Kunimatsu, Nakatsukasa, Sawada, Sakai, Hyodo, Hyodo, Itaya, Nakaya, Saegusa, Mazurier, Saneyoshi, Tsujikawa, Yamamoto &amp; Mbua" authorityYear="2007" box="[625,773,1949,1970]" class="Mammalia" family="Hominidae" genus="Nakalipithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FDF2F814886BF81F" box="[625,773,1949,1970]" italics="true" pageId="5" pageNumber="6">Nakalipithecus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C7D4D72FF9F0F59FCA9FA8988B8FAB8" authorityName="Ishida &amp; Pickford" authorityYear="1997" box="[810,982,1280,1301]" class="Mammalia" genus="Samburupithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FCA9FA8988B8FAB8" box="[810,982,1280,1301]" italics="true" pageId="5" pageNumber="6">Samburupithecus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF9F0F59FB91FA898FD8FAB8" authorityName="Suwa, Kono, Katoh, Asfaw &amp; Beyene" authorityYear="2007" box="[1042,1206,1280,1301]" class="Mammalia" genus="Chororapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FB91FA898FD8FAB8" box="[1042,1206,1280,1301]" italics="true" pageId="5" pageNumber="6">Chororapithecus</emphasis>
</taxonomicName>
, broadly overlap in time with
<emphasis id="B909EAE3FF9F0F59FCEEFA958FD3FA9C" box="[877,1213,1308,1329]" italics="true" pageId="5" pageNumber="6">
<taxonomicName id="4C7D4D72FF9F0F59FCEEFA958F63FA9C" box="[877,1037,1308,1329]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Ouranopithecus</taxonomicName>
,
<taxonomicName id="4C7D4D72FF9F0F59FBA4FA958FD3FA9C" authorityName="von Königswald" authorityYear="1972" box="[1063,1213,1308,1329]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Graecopithecus</taxonomicName>
</emphasis>
and
<taxonomicName id="4C7D4D72FF9F0F59FA93FA958EE0FA9C" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[1296,1422,1308,1329]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FA93FA958EE0FA9C" box="[1296,1422,1308,1329]" italics="true" pageId="5" pageNumber="6">Anadoluvius</emphasis>
</taxonomicName>
<bibRefCitation id="EFEC4B00FF9F0F59FA0DFA9E8ECFFA87" author="Ishida, H. &amp; Pickford, M." box="[1422,1441,1303,1322]" journalOrPublisher="C. R. Acad. Sci. Paris" pageId="5" pageNumber="6" pagination="823 - 829" part="325" refId="ref10430" refString="21. Ishida, H. &amp; Pickford, M. A new late Miocene hominoid from Kenya: Samburupithecus kiptalami gen. et sp. nov. C. R. Acad. Sci. Paris 325, 823 - 829 (1997)." title="A new late Miocene hominoid from Kenya: Samburupithecus kiptalami gen. et sp. nov." type="journal article" year="1997">21</bibRefCitation>
<bibRefCitation id="EFEC4B00FF9F0F59FA2FFA9E8ED1FA87" author="Suwa, G. &amp; Kono, R. T. &amp; Katoh, S. &amp; Berhane, A. &amp; Beyene, Y." box="[1452,1471,1303,1322]" journalOrPublisher="Nature" pageId="5" pageNumber="6" pagination="921 - 924" part="448" refId="ref10523" refString="23. Suwa, G., Kono, R. T., Katoh, S., Berhane, A. &amp; Beyene, Y. A new species of great ape from the late Miocene epoch in Ethiopia. Nature 448, 921 - 924 (2007)." title="A new species of great ape from the late Miocene epoch in Ethiopia" type="journal article" year="2007">23</bibRefCitation>
<taxonomicName id="4C7D4D72FF9F0F59FCA9FABE88D0FAE1" authorityName="Kunimatsu, Nakatsukasa, Sawada, Sakai, Hyodo, Hyodo, Itaya, Nakaya, Saegusa, Mazurier, Saneyoshi, Tsujikawa, Yamamoto &amp; Mbua" authorityYear="2007" box="[810,958,1335,1356]" class="Mammalia" family="Hominidae" genus="Nakalipithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FCA9FABE88D0FAE1" box="[810,958,1335,1356]" italics="true" pageId="5" pageNumber="6">Nakalipithecus</emphasis>
</taxonomicName>
has been identified as potentially ancestral to
<taxonomicName id="4C7D4D72FF9F0F59FCA9FADA88A6FAC5" box="[810,968,1363,1384]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FCA9FADA88A6FAC5" box="[810,968,1363,1384]" italics="true" pageId="5" pageNumber="6">Ouranopithecus</emphasis>
</taxonomicName>
<bibRefCitation id="EFEC4B00FF9F0F59FC4BFAC788B5FACC" author="Kunimatsu, Y. &amp; et al." box="[968,987,1358,1377]" journalOrPublisher="Proc. Natl Acad. Sci. USA" pageId="5" pageNumber="6" pagination="19220 - 19225" part="104" refId="ref10476" refString="22. Kunimatsu, Y. et al. A new Late Miocene great ape from Kenya and its implications for the origins of African great apes and humans. Proc. Natl Acad. Sci. USA 104, 19220 - 19225 (2007)." title="A new Late Miocene great ape from Kenya and its implications for the origins of African great apes and humans" type="journal article" year="2007">22</bibRefCitation>
. However,
<taxonomicName id="4C7D4D72FF9F0F59FBE4FADA8F95FAC5" authorityName="Kunimatsu, Nakatsukasa, Sawada, Sakai, Hyodo, Hyodo, Itaya, Nakaya, Saegusa, Mazurier, Saneyoshi, Tsujikawa, Yamamoto &amp; Mbua" authorityYear="2007" box="[1127,1275,1363,1384]" class="Mammalia" family="Hominidae" genus="Nakalipithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FBE4FADA8F95FAC5" box="[1127,1275,1363,1384]" italics="true" pageId="5" pageNumber="6">Nakalipithecus</emphasis>
</taxonomicName>
differs in many details of dental morphology from
<taxonomicName id="4C7D4D72FF9F0F59FB28FAE68E47FA29" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[1195,1321,1391,1412]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FB28FAE68E47FA29" box="[1195,1321,1391,1412]" italics="true" pageId="5" pageNumber="6">Anadoluvius</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF9F0F59FAEAFAF98815FA0D" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FAEAFAF98815FA0D" italics="true" pageId="5" pageNumber="6">Ouranopithecus</emphasis>
</taxonomicName>
(see Supplementary Note 3). As noted,
<taxonomicName id="4C7D4D72FF9F0F59FAA5FA028ED4FA0D" authorityName="Kunimatsu, Nakatsukasa, Sawada, Sakai, Hyodo, Hyodo, Itaya, Nakaya, Saegusa, Mazurier, Saneyoshi, Tsujikawa, Yamamoto &amp; Mbua" authorityYear="2007" box="[1318,1466,1419,1440]" class="Mammalia" family="Hominidae" genus="Nakalipithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FAA5FA028ED4FA0D" box="[1318,1466,1419,1440]" italics="true" pageId="5" pageNumber="6">Nakalipithecus</emphasis>
</taxonomicName>
, represented in the data matrix by 43% of the total number of characters, consistently falls outside the crown hominids, which fails to support the hypothesis of an ancestral-descendant relationship with
<emphasis id="B909EAE3FF9F0F59FC60FA728F16F9BD" box="[995,1144,1531,1552]" italics="true" pageId="5" pageNumber="6">Ouranopitheus</emphasis>
. It has been hypothesized that both
<taxonomicName id="4C7D4D72FF9F0F59FCEBF99E8F62F981" authorityName="Suwa, Kono, Katoh, Asfaw &amp; Beyene" authorityYear="2007" box="[872,1036,1559,1580]" class="Mammalia" genus="Chororapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FCEBF99E8F62F981" box="[872,1036,1559,1580]" italics="true" pageId="5" pageNumber="6">Chororapithecus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF9F0F59FBD1F99E8F9EF981" box="[1106,1264,1559,1580]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FBD1F99E8F9EF981" box="[1106,1264,1559,1580]" italics="true" pageId="5" pageNumber="6">Ouranopithecus</emphasis>
</taxonomicName>
have phylogenetic affinities with gorillas
<bibRefCitation id="EFEC4B00FF9F0F59FB98F9A78F4BF9EC" author="Dean, D. &amp; Delson, E." box="[1051,1061,1582,1601]" journalOrPublisher="Nature" pageId="5" pageNumber="6" pagination="676 - 677" part="359" refId="ref9913" refString="8. Dean, D. &amp; Delson, E. Second gorilla or third chimp? Nature 359, 676 - 677 (1992)." title="Second gorilla or third chimp?" type="journal article" year="1992">8</bibRefCitation>
,
<bibRefCitation id="EFEC4B00FF9F0F59FBAAF9A78F52F9EC" author="Kunimatsu, Y. &amp; et al." box="[1065,1084,1582,1601]" journalOrPublisher="Proc. Natl Acad. Sci. USA" pageId="5" pageNumber="6" pagination="19220 - 19225" part="104" refId="ref10476" refString="22. Kunimatsu, Y. et al. A new Late Miocene great ape from Kenya and its implications for the origins of African great apes and humans. Proc. Natl Acad. Sci. USA 104, 19220 - 19225 (2007)." title="A new Late Miocene great ape from Kenya and its implications for the origins of African great apes and humans" type="journal article" year="2007">22</bibRefCitation>
,
<bibRefCitation id="EFEC4B00FF9F0F59FBC2F9A78F3AF9EC" author="Suwa, G. &amp; Kono, R. T. &amp; Katoh, S. &amp; Berhane, A. &amp; Beyene, Y." box="[1089,1108,1582,1601]" journalOrPublisher="Nature" pageId="5" pageNumber="6" pagination="921 - 924" part="448" refId="ref10523" refString="23. Suwa, G., Kono, R. T., Katoh, S., Berhane, A. &amp; Beyene, Y. A new species of great ape from the late Miocene epoch in Ethiopia. Nature 448, 921 - 924 (2007)." title="A new species of great ape from the late Miocene epoch in Ethiopia" type="journal article" year="2007">23</bibRefCitation>
,
<bibRefCitation id="EFEC4B00FF9F0F59FBDAF9A78F02F9EC" author="Begun, D. R." box="[1113,1132,1582,1601]" journalOrPublisher="J. Vert. Paleontol." pageId="5" pageNumber="6" pagination="74" part="87" refId="ref10909" refString="33. Begun, D. R. Very old hominoid divergence dates based on paleontological and molecular data. J. Vert. Paleontol. 87, 74 (2015)." title="Very old hominoid divergence dates based on paleontological and molecular data" type="journal article" year="2015">33</bibRefCitation>
. However,
<taxonomicName id="4C7D4D72FF9F0F59FB7BF9BA8EF2F9E5" authorityName="Suwa, Kono, Katoh, Asfaw &amp; Beyene" authorityYear="2007" box="[1272,1436,1587,1608]" class="Mammalia" genus="Chororapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FB7BF9BA8EF2F9E5" box="[1272,1436,1587,1608]" italics="true" pageId="5" pageNumber="6">Chororapithecus</emphasis>
</taxonomicName>
is readily distinguished from both
<taxonomicName id="4C7D4D72FF9F0F59FBECF9C78F83F9CE" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[1135,1261,1614,1635]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FBECF9C78F83F9CE" box="[1135,1261,1614,1635]" italics="true" pageId="5" pageNumber="6">Anadoluvius</emphasis>
</taxonomicName>
and Our
<emphasis id="B909EAE3FF9F0F59FACAF9C78ED1F9CE" box="[1353,1471,1614,1635]" italics="true" pageId="5" pageNumber="6">anopithecus</emphasis>
(see differential diagnosis), and its affinity with gorillas has been questioned
<bibRefCitation id="EFEC4B00FF9F0F59FC14F90888CFF939" author="Begun, D. R." box="[919,929,1665,1684]" editor="Henke, W. &amp; Tattersall, I." journalOrPublisher="Springer" pageId="5" pageNumber="6" pagination="1261 - 1332" refId="ref9660" refString="1. Begun, D. R. Fossil Record of Miocene Hominoids in Handbook of Paleoanthropology (eds. Henke, W. &amp; Tattersall, I.) 1261 - 1332 (Springer, 2015)." title="Fossil Record of Miocene Hominoids" type="book chapter" volumeTitle="Handbook of Paleoanthropology" year="2015">1</bibRefCitation>
,
<bibRefCitation id="EFEC4B00FF9F0F59FC25F90888D7F939" author="Harrison, T." box="[934,953,1665,1684]" editor="Werdelin, L. &amp; Sanders, W. J." journalOrPublisher="University of California Press, Berkeley" pageId="5" pageNumber="6" pagination="429 - 469" refId="ref10942" refString="34. Harrison, T. Dendropithecoidea, Proconsuloidea, and Hominoidea (Catarrhini, Primates) in Cenozoic Mammals of Africa (eds. Werdelin, L. &amp; Sanders, W. J.) 429 - 469 (University of California Press, Berkeley, 2010)." title="Dendropithecoidea, Proconsuloidea, and Hominoidea (Catarrhini, Primates)" type="book chapter" volumeTitle="Cenozoic Mammals of Africa" year="2010">34</bibRefCitation>
A close phylogenetic relationship with gorillas is not supported by our results21. claim that
<taxonomicName id="4C7D4D72FF9F0F59FA90F92B8ED1F91A" authorityName="Ishida &amp; Pickford" authorityYear="1997" box="[1299,1471,1698,1719]" class="Mammalia" genus="Samburupithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FA90F92B8ED1F91A" box="[1299,1471,1698,1719]" italics="true" pageId="5" pageNumber="6">Samburupithecus</emphasis>
</taxonomicName>
has phylogenetic affinities with African apes and humans, though this conclusion has also been challenged by
<bibRefCitation id="EFEC4B00FF9F0F59FB7FF95C8E68F945" author="Begun, D. R." box="[1276,1286,1749,1768]" editor="Henke, W. &amp; Tattersall, I." journalOrPublisher="Springer" pageId="5" pageNumber="6" pagination="1261 - 1332" refId="ref9660" refString="1. Begun, D. R. Fossil Record of Miocene Hominoids in Handbook of Paleoanthropology (eds. Henke, W. &amp; Tattersall, I.) 1261 - 1332 (Springer, 2015)." title="Fossil Record of Miocene Hominoids" type="book chapter" volumeTitle="Handbook of Paleoanthropology" year="2015">1</bibRefCitation>
,
<bibRefCitation id="EFEC4B00FF9F0F59FA89F95C8E73F945" author="Olejniczak, A. J. &amp; Begun, D. R. &amp; Mbua, E. &amp; Hublin, J. J." box="[1290,1309,1749,1768]" journalOrPublisher="Am. J. Phys. Anthropol." pageId="5" pageNumber="6" pagination="202" part="48" refId="ref10993" refString="35. Olejniczak, A. J., Begun, D. R., Mbua, E. &amp; Hublin, J. J. New evidence on the phylogenetic position of Samburupithecus. Am. J. Phys. Anthropol. 48, 202 (2009)." title="New evidence on the phylogenetic position of Samburupithecus" type="journal article" year="2009">35</bibRefCitation>
, who conclude that
<taxonomicName id="4C7D4D72FF9F0F59FCD4F97F8F6DF8A6" authorityName="Ishida &amp; Pickford" authorityYear="1997" box="[855,1027,1782,1803]" class="Mammalia" genus="Samburupithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FCD4F97F8F6DF8A6" box="[855,1027,1782,1803]" italics="true" pageId="5" pageNumber="6">Samburupithecus</emphasis>
</taxonomicName>
is not a hominid but instead a vestige of the early Miocene proconsuloid radiation. As with
<taxonomicName id="4C7D4D72FF9F0F59FA98F89B8ED1F88A" authorityName="Suwa, Kono, Katoh, Asfaw &amp; Beyene" authorityYear="2007" box="[1307,1471,1810,1831]" class="Mammalia" genus="Chororapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9F0F59FA98F89B8ED1F88A" box="[1307,1471,1810,1831]" italics="true" pageId="5" pageNumber="6">Chororapithecus</emphasis>
</taxonomicName>
and
<emphasis id="B909EAE3FF9F0F59FCDDF8A78FDDF8EE" box="[862,1203,1838,1859]" italics="true" pageId="5" pageNumber="6">
<taxonomicName id="4C7D4D72FF9F0F59FCDDF8A7889AF8EE" authorityName="Kunimatsu, Nakatsukasa, Sawada, Sakai, Hyodo, Hyodo, Itaya, Nakaya, Saegusa, Mazurier, Saneyoshi, Tsujikawa, Yamamoto &amp; Mbua" authorityYear="2007" box="[862,1012,1838,1859]" class="Mammalia" family="Hominidae" genus="Nakalipithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Nakalipithecus</taxonomicName>
,
<taxonomicName id="4C7D4D72FF9F0F59FB84F8A78FDDF8EE" authorityName="Ishida &amp; Pickford" authorityYear="1997" box="[1031,1203,1838,1859]" class="Mammalia" genus="Samburupithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Samburupithecus</taxonomicName>
</emphasis>
falls outside the crown hominid clade.
</paragraph>
</subSubSection>
<subSubSection id="C367657AFF9F0F5AFCA9F80B8E7CFE84" lastPageId="6" lastPageNumber="7" pageId="5" pageNumber="6" type="biology_ecology">
<paragraph id="8BC236F1FF9F0F5AFCA9F80B8B95FC4E" blockId="5.[810,1472,1921,1971]" lastBlockId="6.[116,779,136,1972]" lastPageId="6" lastPageNumber="7" pageId="5" pageNumber="6">
<emphasis id="B909EAE3FF9F0F59FCA9F80B8F1EF83B" box="[810,1136,1921,1943]" italics="true" pageId="5" pageNumber="6">Diversity and paleobiogeography</emphasis>
. A comprehensive review of the taxonomy and phylogeny of late Miocene apes is needed, given recent discoveries and reinterpretations. Here we focus on the diversity and paleobiogeographic implications of
<taxonomicName id="4C7D4D72FF9C0F5AFD0FFF2D8864FF14" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[652,778,164,185]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFD0FFF2D8864FF14" box="[652,778,164,185]" italics="true" pageId="6" pageNumber="7">Anadoluvius</emphasis>
</taxonomicName>
(Supplementary Fig. 15; Supplementary Table 8).
<taxonomicName id="4C7D4D72FF9C0F5AFD0FFF498864FF78" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[652,778,192,213]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFD0FFF498864FF78" box="[652,778,192,213]" italics="true" pageId="6" pageNumber="7">Anadoluvius</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF9C0F5AFF22FF558A51FF5C" box="[161,319,220,241]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFF22FF558A51FF5C" box="[161,319,220,241]" italics="true" pageId="6" pageNumber="7">Ouranopithecus</emphasis>
</taxonomicName>
share attributes with other European middle and late Miocene hominids that distinguish them from late Miocene ape fossils from Africa and the broadly contemporaneous pongines
<emphasis id="B909EAE3FF9C0F5AFE13FEB989ABFEE8" box="[400,709,304,325]" italics="true" pageId="6" pageNumber="7">
<taxonomicName id="4C7D4D72FF9C0F5AFE13FEB98963FEE8" box="[400,525,304,325]" class="Mammalia" family="Hominidae" genus="Sivapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">Sivapithecus</taxonomicName>
,
<taxonomicName id="4C7D4D72FF9C0F5AFDAAFEB989ABFEE8" box="[553,709,304,325]" class="Mammalia" family="Hominidae" genus="Ankarapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">Ankarapithecus</taxonomicName>
</emphasis>
, and
<taxonomicName id="4C7D4D72FF9C0F5AFFF7FEC58A65FECC" authorityName="Chaimanee, Suteethorn, Jintasakul, Vidthayanon, Marandat &amp; Jaeger" authorityYear="2004" box="[116,267,332,353]" class="Mammalia" family="Hominidae" genus="Khoratpithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFFF7FEC58A65FECC" box="[116,267,332,353]" italics="true" pageId="6" pageNumber="7">Khoratpithecus</emphasis>
</taxonomicName>
(Supplementary Note 3).
<taxonomicName id="4C7D4D72FF9C0F5AFDAFFEC589A4FECC" box="[556,714,332,353]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFDAFFEC589A4FECC" box="[556,714,332,353]" italics="true" pageId="6" pageNumber="7">Ouranopithecus</emphasis>
</taxonomicName>
from Ravin de la Pluie and Xirochori are dated to 9.6 and 9.3 Ma respectively
<superScript id="7C089BB9FF9C0F5AFF6AFEF78A6FFE3C" attach="left" box="[233,257,382,401]" fontSize="8" pageId="6" pageNumber="7">
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,
<bibRefCitation id="EFEC4B00FF9C0F5AFF74FEF78A6FFE3C" author="Koufos, G. D." box="[247,257,382,401]" journalOrPublisher="Geobios" pageId="6" pageNumber="7" pagination="3 - 10" part="49" refId="ref9888" refString="7. Koufos, G. D. History, stratigraphy and fossiliferous sites. Geobios 49, 3 - 10 (2016)." title="History, stratigraphy and fossiliferous sites" type="journal article" year="2016">7</bibRefCitation>
</superScript>
.
<taxonomicName id="4C7D4D72FF9C0F5AFE8EFE0A8AC5FE35" box="[269,427,387,408]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFE8EFE0A8AC5FE35" box="[269,427,387,408]" italics="true" pageId="6" pageNumber="7">Ouranopithecus</emphasis>
</taxonomicName>
from Nikiti 1 is dated to 8.9 Ma
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,
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. Çorakyerler is dated to 8.7 Ma (Supplementary Figs. 16, 17; Supplementary Table 9 and Supplementary Note 6). The Nikiti mandible and maxilla are distinct from
<taxonomicName id="4C7D4D72FF9C0F5AFD88FE5E89C7FE41" box="[523,681,471,492]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFD88FE5E89C7FE41" box="[523,681,471,492]" italics="true" pageId="6" pageNumber="7">Ouranopithecus</emphasis>
</taxonomicName>
from the more northern Macedonian sites and may represent a different taxon
<bibRefCitation id="EFEC4B00FF9C0F5AFF2EFD838BAEFDB0" author="Begun, D. R. &amp; Bohme, M. &amp; Spassov, N. &amp; Sevim Erol, A. &amp; Yenner Yavuz, A." box="[173,192,522,541]" journalOrPublisher="Am. J. Phys. Anthr." pageId="6" pageNumber="7" pagination="15" part="168" refId="ref10108" refString="13. Begun, D. R., Bohme, M., Spassov, N., Sevim Erol, A. &amp; Yenner Yavuz, A. Hominin origins: New evidence from the eastern Mediterranean. Am. J. Phys. Anthr. 168, 15 (2019)." title="Hominin origins: New evidence from the eastern Mediterranean" type="journal article" year="2019">
<superScript id="7C089BB9FF9C0F5AFF2EFD838BAEFDB0" attach="left" box="[173,192,522,541]" fontSize="8" pageId="6" pageNumber="7">13</superScript>
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. This possibility, which needs further study, is interesting in terms of regional evolution as Nikiti 1 is likely to be slightly older than Çorakyerler
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,
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,
<bibRefCitation id="EFEC4B00FF9C0F5AFE05FDCB8AF7FDF8" author="Sevim Erol, A. &amp; et al." box="[390,409,578,597]" pageId="6" pageNumber="7" refId="ref10749" refString="29. Sevim Erol, A. et al. The late Miocene (early Turolian, MN 11) fauna of Corakyerler (Turkey): New collection of large mammals (2017)." title="The late Miocene (early Turolian, MN 11) fauna of Corakyerler (Turkey): New collection of large mammals" type="book" year="2017">29</bibRefCitation>
</superScript>
. Nikiti,
<taxonomicName id="4C7D4D72FF9C0F5AFE78FDCE8917FDF1" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[507,633,583,604]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFE78FDCE8917FDF1" box="[507,633,583,604]" italics="true" pageId="6" pageNumber="7">Anadoluvius</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF9C0F5AFD3FFDCE8BABFDD5" authorityName="von Königswald" authorityYear="1972" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFD3FFDCE8BABFDD5" italics="true" pageId="6" pageNumber="7">Graecopithecus</emphasis>
</taxonomicName>
are distinguished from
<taxonomicName id="4C7D4D72FF9C0F5AFE3EFDEA8935FDD5" box="[445,603,611,632]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFE3EFDEA8935FDD5" box="[445,603,611,632]" italics="true" pageId="6" pageNumber="7">Ouranopithecus</emphasis>
</taxonomicName>
in our PCA and in having a greater degree of canine reduction, elongated P
<subScript id="17F934B4FF9C0F5AFD50FD0189B2FD3B" attach="left" box="[723,732,648,662]" fontSize="6" pageId="6" pageNumber="7">4</subScript>
and large molars relative to mandibular corpus size.
<taxonomicName id="4C7D4D72FF9C0F5AFD0FFD138864FD02" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[652,778,666,687]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFD0FFD138864FD02" box="[652,778,666,687]" italics="true" pageId="6" pageNumber="7">Anadoluvius</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF9C0F5AFF24FD3F8A53FD66" authorityName="von Königswald" authorityYear="1972" box="[167,317,694,715]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFF24FD3F8A53FD66" box="[167,317,694,715]" italics="true" pageId="6" pageNumber="7">Graecopithecus</emphasis>
</taxonomicName>
are distinguished from
<taxonomicName id="4C7D4D72FF9C0F5AFDC9FD3F8986FD66" box="[586,744,694,715]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFDC9FD3F8986FD66" box="[586,744,694,715]" italics="true" pageId="6" pageNumber="7">Ouranopithecus</emphasis>
</taxonomicName>
in root morphology (unknown in NKT 21.) NKT 21 and
<taxonomicName id="4C7D4D72FF9C0F5AFD3FFD5A8BABFCAE" authorityName="von Königswald" authorityYear="1972" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFD3FFD5A8BABFCAE" italics="true" pageId="6" pageNumber="7">Graecopithecus</emphasis>
</taxonomicName>
have inferior transverse tori positioned posterior to the mesial edge of the M
<subScript id="17F934B4FF9C0F5AFEC4FC9D8A3EFC8F" attach="left" box="[327,336,788,802]" fontSize="6" pageId="6" pageNumber="7">1</subScript>
(unknown in
<taxonomicName id="4C7D4D72FF9C0F5AFE62FC838931FCB2" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[481,607,778,799]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFE62FC838931FCB2" box="[481,607,778,799]" italics="true" pageId="6" pageNumber="7">Anadoluvius</emphasis>
</taxonomicName>
), while the torus is anterior to the M
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in
<emphasis id="B909EAE3FF9C0F5AFEE4FCAF89C6FC96" box="[359,680,806,827]" italics="true" pageId="6" pageNumber="7">
<taxonomicName id="4C7D4D72FF9C0F5AFEE4FCAF8966FC96" box="[359,520,806,827]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">Ouranopithecus</taxonomicName>
.
<taxonomicName id="4C7D4D72FF9C0F5AFD91FCAF89C6FC96" authorityName="von Königswald" authorityYear="1972" box="[530,680,806,827]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">Graecopithecus</taxonomicName>
</emphasis>
, which is considerably younger (~7.2 Ma)
<bibRefCitation id="EFEC4B00FF9C0F5AFE39FCB48AAAFCFD" author="Bohme, M. &amp; et al." box="[442,452,829,848]" journalOrPublisher="PLoS ONE" pageId="6" pageNumber="7" pagination="0177347" part="12" refId="ref9782" refString="4. Bohme, M. et al. Messinian age and savannah environment of the possible hominin Graecopithecus from Europe. PLoS ONE 12, e 0177347 (2017)." title="Messinian age and savannah environment of the possible hominin Graecopithecus from Europe" type="journal article" year="2017">
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, has been shown to differ from
<taxonomicName id="4C7D4D72FF9C0F5AFFF7FCD78A7CFCDE" box="[116,274,862,883]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFFF7FCD78A7CFCDE" box="[116,274,862,883]" italics="true" pageId="6" pageNumber="7">Ouranopithecus</emphasis>
</taxonomicName>
in morphology that replicates differences between late Miocene apes and early hominins, such as reduced relative canine size and premolar root morphology
<bibRefCitation id="EFEC4B00FF9C0F5AFD0FFC1889F1FC09" author="Fuss, J. &amp; Spassov, N. &amp; Begun, D. R. &amp; Bohme, M." box="[652,671,913,932]" journalOrPublisher="PLoS ONE" pageId="6" pageNumber="7" pagination="0177127" part="12" refId="ref10011" refString="11. Fuss, J., Spassov, N., Begun, D. R. &amp; Bohme, M. Potential hominin affinities of Graecopithecus from the Late Miocene of Europe. PLoS ONE 12, e 0177127 (2017)." title="Potential hominin affinities of Graecopithecus from the Late Miocene of Europe" type="journal article" year="2017">
<superScript id="7C089BB9FF9C0F5AFD0FFC1889F1FC09" attach="left" box="[652,671,913,932]" fontSize="8" pageId="6" pageNumber="7">11</superScript>
</bibRefCitation>
However, the Nikiti specimens have not previously been included in these comparisons.
</paragraph>
<paragraph id="8BC236F1FF9C0F5AFF0DFC608BB5F915" blockId="6.[116,779,136,1972]" pageId="6" pageNumber="7">
Quantitative and qualitative comparisons reveal many differences among Balkan/Anatolian taxa, indicative of a greater diversity of late Miocene eastern Mediterranean hominids than previously recognized. Hominines appear to have been present and diverse for millions of years in the late Miocene of Europe. Based on the large number of qualitative and quantitative differences among the samples from Macedonia, Attica and Anatolia, we conclude that they represent at least three hominine genera,
<taxonomicName id="4C7D4D72FF9C0F5AFF4BFB418A08FB70" box="[200,358,1224,1245]" class="Mammalia" family="Hominidae" genus="Ouranopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFF4BFB418A08FB70" box="[200,358,1224,1245]" italics="true" pageId="6" pageNumber="7">Ouranopithecus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C7D4D72FF9C0F5AFEFAFB418961FB70" authorityName="von Königswald" authorityYear="1972" box="[377,527,1224,1245]" class="Mammalia" family="Hominidae" genus="Graecopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFEFAFB418961FB70" box="[377,527,1224,1245]" italics="true" pageId="6" pageNumber="7">Graecopithecus</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C7D4D72FF9C0F5AFDD0FB4189BFFB70" authorityName="Sevim-Erol &amp; Begun &amp; Sözer &amp; Mayda &amp; van den Hoek Ostende &amp; Martin &amp; Alçiçek" authorityYear="2023" box="[595,721,1224,1245]" class="Mammalia" family="Hominidae" genus="Anadoluvius" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFDD0FB4189BFFB70" box="[595,721,1224,1245]" italics="true" pageId="6" pageNumber="7">Anadoluvius</emphasis>
</taxonomicName>
. The diversity of hominines in the eastern Mediterranean mirrors that among australopithecines in the Plio-Pleistocene hominin record in Africa. In the phylogenetic analysis presented here the Balkan/ Anatolian taxa are in the sister clade of crown hominines. The fact that dryopithecins are also classified as hominines in this analysis suggests that there was an in situ evolution of thickly enameled late Miocene eastern Mediterranean hominines from more thinly enameled precursors in central and western Europe, though this conclusion has been challenged
<bibRefCitation id="EFEC4B00FF9C0F5AFDBFFA378928FA7C" author="Pugh, K. D." box="[572,582,1470,1489]" journalOrPublisher="J. Hum. Evol" pageId="6" pageNumber="7" refId="ref9940" refString="9. Pugh, K. D. Phylogenetic analysis of Middle-Late Miocene apes. J. Hum. Evol. https: // doi. org / 10.1016 / j. jhevol. 2021.103140 (2022)." title="Phylogenetic analysis of Middle-Late Miocene apes" type="book" year="2022">9</bibRefCitation>
,
<bibRefCitation id="EFEC4B00FF9C0F5AFDC9FA378933FA7C" author="Alba, D. M." box="[586,605,1470,1489]" journalOrPublisher="Evol. Anthropol." pageId="6" pageNumber="7" pagination="254 - 269" part="21" refId="ref10575" refString="24. Alba, D. M. Fossil apes from the Valles-Penedes basin. Evol. Anthropol. 21, 254 - 269 (2012)." title="Fossil apes from the Valles-Penedes basin" type="journal article" year="2012">24</bibRefCitation>
.
<emphasis id="B909EAE3FF9C0F5AFDEEFA4A8927FA59" italics="true" pageId="6" pageNumber="7">
<taxonomicName id="4C7D4D72FF9C0F5AFDEEFA4A8A6AFA59" authority=", Anoiapithecus" authorityName="Anoiapithecus" class="Mammalia" family="Hominidae" genus="Pierolapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">Pierolapithecus, Anoiapithecus</taxonomicName>
,
<taxonomicName id="4C7D4D72FF9C0F5AFE91FA568AF8FA59" authorityName="Lartet" authorityYear="1856" box="[274,406,1503,1524]" class="Mammalia" family="Hominidae" genus="Dryopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">Dryopithecus</taxonomicName>
,
<taxonomicName id="4C7D4D72FF9C0F5AFE26FA568927FA59" authorityName="Villalta &amp; Crusafont Pairó" authorityYear="1944" box="[421,585,1503,1524]" class="Mammalia" family="Hominidae" genus="Hispanopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">Hispanopithecus</taxonomicName>
</emphasis>
and
<taxonomicName id="4C7D4D72FF9C0F5AFD07FA568864FA59" authorityName="Kretzoi" authorityYear="1969" box="[644,778,1503,1524]" class="Mammalia" family="Hominidae" genus="Rudapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFD07FA568864FA59" box="[644,778,1503,1524]" italics="true" pageId="6" pageNumber="7">Rudapithecus</emphasis>
</taxonomicName>
all share attributes with extant hominines and are distinguished from pongines such as
<taxonomicName id="4C7D4D72FF9C0F5AFED8F99E8ABBF981" box="[347,469,1559,1580]" class="Mammalia" family="Hominidae" genus="Sivapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFED8F99E8ABBF981" box="[347,469,1559,1580]" italics="true" pageId="6" pageNumber="7">Sivapithecus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7D4D72FF9C0F5AFD85F99E89CCF981" box="[518,674,1559,1580]" class="Mammalia" family="Hominidae" genus="Ankarapithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFD85F99E89CCF981" box="[518,674,1559,1580]" italics="true" pageId="6" pageNumber="7">Ankarapithecus</emphasis>
</taxonomicName>
(
<tableCitation id="C6FF034AFF9C0F5AFD2CF99E886AF981" box="[687,772,1559,1581]" captionStart="Table 1" captionStartId="1.[127,184,1631,1653]" captionTargetBox="[127,767,1722,1956]" captionTargetId="graphics-713@1.[116,777,1609,1966]" captionText="Table 1 Hominine and pongine synapomorphies (18 taxa, unordered)." httpUri="http://table.plazi.org/id/DF026679FF9B0F5DFFFCF9D68B97F93F" pageId="6" pageNumber="7" tableUuid="DF026679FF9B0F5DFFFCF9D68B97F93F">Tables 1</tableCitation>
,
<tableCitation id="C6FF034AFF9C0F5AFFF7F9BA8BEEF9E5" box="[116,128,1587,1608]" captionStart="Table 2" captionStartId="1.[820,877,1053,1075]" captionTargetBox="[820,1460,1145,1482]" captionTargetId="graphics-777@1.[810,1472,1031,1491]" captionText="Table 2 Hominine and pongine synapomorphies (18 taxa, ordered)." httpUri="http://table.plazi.org/id/DF026679FF9B0F5DFCB7FB9488FBFBFD" pageId="6" pageNumber="7" tableUuid="DF026679FF9B0F5DFCB7FB9488FBFBFD">2</tableCitation>
)
<bibRefCitation id="EFEC4B00FF9C0F5AFF0AF9A78BFDF9EC" author="Begun, D. R." box="[137,147,1582,1601]" editor="Henke, W. &amp; Tattersall, I." journalOrPublisher="Springer" pageId="6" pageNumber="7" pagination="1261 - 1332" refId="ref9660" refString="1. Begun, D. R. Fossil Record of Miocene Hominoids in Handbook of Paleoanthropology (eds. Henke, W. &amp; Tattersall, I.) 1261 - 1332 (Springer, 2015)." title="Fossil Record of Miocene Hominoids" type="book chapter" volumeTitle="Handbook of Paleoanthropology" year="2015">1</bibRefCitation>
<bibRefCitation id="EFEC4B00FF9C0F5AFF1EF9A78BC9F9EC" author="Begun, D. R." box="[157,167,1582,1601]" journalOrPublisher="Geodiversitas" pageId="6" pageNumber="7" pagination="789 - 816" part="31" refId="ref9746" refString="3. Begun, D. R. Dryopithecus, Darwin, de Bonis and the European origin of the African apes and human clade. Geodiversitas 31, 789 - 816 (2009)." title="Dryopithecus, Darwin, de Bonis and the European origin of the African apes and human clade" type="journal article" year="2009">3</bibRefCitation>
,
<bibRefCitation id="EFEC4B00FF9C0F5AFF28F9A78BD0F9EC" author="Begun, D. R." box="[171,190,1582,1601]" editor="Hartwig, W." journalOrPublisher="Cambridge University Press" pageId="6" pageNumber="7" pagination="339 - 368" refId="ref11044" refString="36. Begun, D. R. European hominoids in The Primate Fossil Record (ed. Hartwig, W.) 339 - 368 (Cambridge University Press, 2002)." title="European hominoids" type="book chapter" volumeTitle="The Primate Fossil Record" year="2002">36</bibRefCitation>
,
<bibRefCitation id="EFEC4B00FF9C0F5AFF40F9A78BB8F9EC" author="Kelley, J." box="[195,214,1582,1601]" editor="Hartwig, W." journalOrPublisher="Cambridge University Press" pageId="6" pageNumber="7" pagination="369 - 384" refId="ref11078" refString="37. Kelley, J. The hominoid radiation in Asia, in The Primate Fossil Record (ed. Hartwig, W.) 369 - 384 (Cambridge University Press, 2002)." title="The hominoid radiation in Asia" type="book chapter" volumeTitle="The Primate Fossil Record" year="2002">37</bibRefCitation>
, (but see refs.
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,
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for alternative views). A clade that includes both thinly and thickly enameled taxa, in this case the dryopithecins and the Balkan/Anatolian apes, has a parallel in Africa with
<taxonomicName id="4C7D4D72FF9C0F5AFF68F90E8A07F931" box="[235,361,1671,1692]" class="Mammalia" family="Hominidae" genus="Ardipithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFF68F90E8A07F931" box="[235,361,1671,1692]" italics="true" pageId="6" pageNumber="7">Ardipithecus</emphasis>
</taxonomicName>
and australopithecines and with
<taxonomicName id="4C7D4D72FF9C0F5AFD35F90E89B0F931" authorityName="Oken" authorityYear="1816" box="[694,734,1671,1692]" class="Mammalia" family="Hominidae" genus="Pan" kingdom="Animalia" order="Primates" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B909EAE3FF9C0F5AFD35F90E89B0F931" box="[694,734,1671,1692]" italics="true" pageId="6" pageNumber="7">Pan</emphasis>
</taxonomicName>
and hominins.
</paragraph>
<paragraph id="8BC236F1FF9C0F5AFF0DF9378E7CFE84" blockId="6.[116,779,136,1972]" lastBlockId="6.[810,1472,136,298]" pageId="6" pageNumber="7">
The parallel evolution of thickly and thinly enameled members of a clade in Africa and Europe is not proof that the late Miocene European apes are all hominines, but it does make this hypothesis, supported by the results of our cladistic analysis, plausible. It is possible that the generally more thinly enameled dryopithecins and the later occurring thickly enameled Balkan/ Anatolian hominines do not share an ancestor-descendant relationship and represent separate dispersal events into Europe from Africa (e.g. ref.
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), though this is less parsimonious biogeographically and contrasts with the results of the phylogenetic analysis presented here. While independent dispersal events are possible, we regard the in situ European hypothesis as more likely and more parsimonious given the current evidence. Other independent lines of evidence are also consistent with the widespread presence of hominines in Europe
<bibRefCitation id="EFEC4B00FF9C0F5AFB7BFE868E65FE8F" author="Johnson, R. J. &amp; Andrews, P." box="[1272,1291,271,290]" journalOrPublisher="Evol. Anthropol." pageId="6" pageNumber="7" pagination="250 - 257" part="19" refId="ref11114" refString="38. Johnson, R. J. &amp; Andrews, P. Fructose, uricase, and the Back-to-Africa hypothesis. Evol. Anthropol. 19, 250 - 257 (2010)." title="Fructose, uricase, and the Back-to-Africa hypothesis" type="journal article" year="2010">38</bibRefCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>