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<document id="7CFE8CAA202B7318AE70CD8EF814647C" ID-CLB-Dataset="37519" ID-DOI="10.1126/science.aab2625" ID-GBIF-Dataset="15211f60-f22d-48f0-8c7a-1bb8c31acee1" ID-Zenodo-Dep="269424" ID-ZooBank="lsid:zoobank.org:pub:4A5EC1F1-29BD-4925-8CC6-04AB083C61DA" IM.metadata_requiresApprovalFor="plazi" IM.taxonomicNames_requiresApprovalFor="plazi" checkinTime="1464163785068" checkinUser="plazi" docAuthor="David M. Alba, Sergio Almécija, Daniel DeMiguel, Josep Fortuny, Miriam Pérez de los Ríos, Marta Pina, Josep M. Robles &amp; Salvador Moyà-Solà" docDate="2015" docId="924187F6FA7E8A1D5A75FF53FC1AA479" docLanguage="en" docName="aab2625.full.pdf" docOrigin="Science 350 (6260)" docStyle="DocumentStyle{}" docTitle="Pliobates cataloniae Alba, Almécija, DeMiguel, Fortuny, Ríos, Pina, Robles &amp; Moyà-Solà, 2015, gen. et sp. nov." docType="treatment" docUuid="3FB80ECA-174A-4D5E-9699-E5C960F2A17B" docUuidSource="ZooBank" docVersion="12" lastPageNumber="8" masterDocId="6E78FF8EFA7A8A155848FFBAFFA9A57C" masterDocTitle="Miocene small-bodied ape from Eurasia sheds light on hominoid evolution" masterLastPageNumber="11" masterPageNumber="1" pageNumber="4" updateTime="1698662623451" updateUser="ExternalLinkService">
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<mods:title id="C9F847A979745D43AE796EED2A48A6B0">Miocene small-bodied ape from Eurasia sheds light on hominoid evolution</mods:title>
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<mods:namePart id="8F91A41620A2375AFBB52E0F011DC353">David M. Alba</mods:namePart>
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<mods:namePart id="DF066D7F855CFE52D45AA630D93534C6">Sergio Almécija</mods:namePart>
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<mods:namePart id="7C0B0C15538807A118FC5BDAC2517B78">Daniel DeMiguel</mods:namePart>
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<mods:namePart id="9803B44B71E975A9F97090BB769CB1B2">Josep Fortuny</mods:namePart>
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<mods:namePart id="CA6867F6099B02A8C9DAF4EB1FC3D3AA">Miriam Pérez de los Ríos</mods:namePart>
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<mods:namePart id="86313072D28A2CAB24B7212EB10C7899">Marta Pina</mods:namePart>
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<mods:namePart id="8C4511BE24F04283D9E45A3356727CC5">Josep M. Robles</mods:namePart>
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<mods:namePart id="DADB36A44FCBD1B02AAB1CD878C01C9A">Salvador Moyà-Solà</mods:namePart>
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<treatment id="924187F6FA7E8A1D5A75FF53FC1AA479" ID-DOI="http://doi.org/10.5281/zenodo.4420189" ID-GBIF-Taxon="125135179" ID-Zenodo-Dep="4420189" LSID="urn:lsid:zoobank.org:act:3FB80ECA-174A-4D5E-9699-E5C960F2A17B" httpUri="http://treatment.plazi.org/id/924187F6FA7E8A1D5A75FF53FC1AA479" lastPageId="8" lastPageNumber="8" pageId="4" pageNumber="4">
<subSubSection id="52F2656BFA7E8A115A75FF53FC17A581" box="[573,958,233,253]" pageId="4" pageNumber="4" type="nomenclature">
<paragraph id="1A5736E0FA7E8A115A75FF53FC17A581" blockId="4.[573,1012,233,1323]" box="[573,958,233,253]" pageId="4" pageNumber="4">
<heading id="411F818CFA7E8A115A75FF53FC17A581" bold="true" box="[573,958,233,253]" fontSize="8" level="8" pageId="4" pageNumber="4" reason="0">
<emphasis id="289CEAF2FA7E8A115A75FF53FC17A581" bold="true" box="[573,958,233,253]" pageId="4" pageNumber="4">
<taxonomicName id="DDE84D63FA7E8A115A75FF53FCBEA580" box="[573,791,233,252]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="4" pageNumber="4" phylum="Chordata" rank="species" species="cataloniae" status="gen. et sp. nov.">Pliobates cataloniae</taxonomicName>
<taxonomicNameLabel id="33AF5789FA7E8A115B55FF50FC17A581" box="[797,958,234,253]" pageId="4" pageNumber="4">
<emphasis id="289CEAF2FA7E8A115B55FF50FC17A581" bold="true" box="[797,958,234,253]" italics="true" pageId="4" pageNumber="4">gen. et sp. nov.</emphasis>
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</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="52F2656BFA7E8A115A75FEB6FD2DA6AE" pageId="4" pageNumber="4" type="materials_examined">
<paragraph id="1A5736E0FA7E8A115A75FEB6FD14A7EF" blockId="4.[573,1012,233,1323]" pageId="4" pageNumber="4">
Holotype: IPS58443, a partial skeleton with an associated skull (
<figureCitation id="82D32A65FA7E8A115A88FE9DFD47A446" box="[704,750,295,314]" captionStart="Fig. 1" captionStartId="2.[98,129,1811,1828]" captionTargetBox="[115,993,753,1778]" captionTargetId="figure@2.[115,993,754,1780]" captionTargetPageId="2" captionText="Fig. 1. Cranium and dentition. (A to C) Cranium of the holotype (IPS 58443) of Pliobates cataloniae gen. et sp. nov. The main cranial fragments, including the basicranium and the right palate, are shown in basal view (A); details of the right palatal fragment are shown in left-lateral (B) and right-lateral (C) views. (D) Detail of the right postcanine teeth, in occlusal view (mesial is to the right)." httpUri="https://zenodo.org/record/269425/files/figure.png" pageId="4" pageNumber="4">Fig. 1</figureCitation>
and movie S1), housed at the ICP. It is composed of 70 bones and bone fragments (table S1) found in close spatial association, which, given the lack of repeated elements, are attributed to a single adult female individual (based on the small canine alveolus), with an estimated body mass of 4 to 5 kg (tables S7 and S8). It includes large portions of the cranium with postcanine maxillary teeth (
<tableCitation id="576A035BFA7E8A115B2CFE41FC0DA772" box="[868,932,507,526]" captionStart="Table 1" captionStartId="4.[605,653,1497,1513]" captionText="Table 1. Dental measurements. Standard dental measurements to assess dental size and proportions were taken to the nearest 0.1 mm in the holotype (IPS 58443) of Pliobates cataloniae gen. et sp. nov. MD, mesiodistal length (in millimeters); BL, buccolingual width (in millimeters); BLI, breadth / length index (in percent), computed as BL / MD × 100. Dashes indicate lack of data due to incomplete preservation." pageId="4" pageNumber="4">Table 1</tableCitation>
), a mandibular fragment, a partial left forelimb (nearly complete humerus, radius, partial ulna, carpals, and bones of the manual rays), more fragmentary elements of the right forelimb, and bones from the hind limb.
</paragraph>
<paragraph id="1A5736E0FA7E8A115A1AFD21FD0AA781" blockId="4.[573,1012,233,1323]" pageId="4" pageNumber="4">Type locality: ACM/C8-A4 (els Hostalets de Pierola, Catalonia, Spain), in the ACM stratigraphic series (Vallès-Penedès Basin, northeast Iberian Peninsula).</paragraph>
<paragraph id="1A5736E0FA7E8A115A1AFCBFFD2DA6AE" blockId="4.[573,1012,233,1323]" pageId="4" pageNumber="4">
Age, stratigraphic position, and distribution: Only known from the type locality, which has an estimated age of 11.6 Ma (middle/late Miocene boundary) and is thus somewhat younger than all other ACM hominoid- and pliopithecoid-bearing localities (
<emphasis id="289CEAF2FA7E8A115AD4FC30FD0EA6E1" bold="true" box="[668,679,906,925]" italics="true" pageId="4" pageNumber="4">9</emphasis>
,
<emphasis id="289CEAF2FA7E8A115AFDFC30FD61A6E1" bold="true" box="[693,712,906,925]" italics="true" pageId="4" pageNumber="4">31</emphasis>
,
<emphasis id="289CEAF2FA7E8A115A9EFC30FD42A6E1" bold="true" box="[726,747,906,925]" italics="true" pageId="4" pageNumber="4">33</emphasis>
,
<emphasis id="289CEAF2FA7E8A115AB1FC30FCA2A6E1" bold="true" box="[761,779,906,925]" italics="true" pageId="4" pageNumber="4">77</emphasis>
,
<emphasis id="289CEAF2FA7E8A115B51FC30FC84A6E1" bold="true" box="[793,813,906,925]" italics="true" pageId="4" pageNumber="4">78</emphasis>
), the latter of which have been dated to 11.7 to 11.9 Ma [updated from (
<emphasis id="289CEAF2FA7E8A115A0BFC05FDFDA6AE" bold="true" box="[579,596,959,978]" italics="true" pageId="4" pageNumber="4">77</emphasis>
,
<emphasis id="289CEAF2FA7E8A115A17FC05FDDBA6AE" bold="true" box="[607,626,959,978]" italics="true" pageId="4" pageNumber="4">78</emphasis>
)].
</paragraph>
</subSubSection>
<subSubSection id="52F2656BFA7E8A115A1AFC60FC98A057" pageId="4" pageNumber="4" type="etymology">
<paragraph id="1A5736E0FA7E8A115A1AFC60FC98A057" blockId="4.[573,1012,233,1323]" pageId="4" pageNumber="4">
Etymology: Genus name from the Latin
<emphasis id="289CEAF2FA7E8A115B82FC60FC5AA691" bold="true" box="[970,1011,986,1005]" italics="true" pageId="4" pageNumber="4">plio-</emphasis>
(itself from the Greek, meaning “greater in extent”) and from the Greek
<emphasis id="289CEAF2FA7E8A115B71FBB5FCCEA15E" bold="true" box="[825,871,1039,1058]" italics="true" pageId="4" pageNumber="4">bates</emphasis>
(meaning “the one that walks or haunts”). The name is a contraction of the genus names
<emphasis id="289CEAF2FA7E8A115B09FBFEFC07A12B" bold="true" box="[833,942,1092,1111]" italics="true" pageId="4" pageNumber="4">Pliopithecus</emphasis>
(“more ape”) and
<taxonomicName id="DDE84D63FA7E8A115AD4FBE4FD5FA10D" box="[668,758,1118,1137]" class="Mammalia" family="Hylobatidae" genus="Hylobates" kingdom="Animalia" order="Primates" pageId="4" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="289CEAF2FA7E8A115AD4FBE4FD5FA10D" bold="true" box="[668,758,1118,1137]" italics="true" pageId="4" pageNumber="4">Hylobates</emphasis>
</taxonomicName>
(“the one that walks in the woods or in the trees”), in allusion to the small body size and the mosaic of primitive (stem catarrhinelike) and derived (crown hominoid) features displayed by the new taxon. The species epithet is the genitive of the female substantive “Catalonia,” the Latin name of Catalunya (in which the type locality is situated).
</paragraph>
</subSubSection>
<subSubSection id="52F2656BFA7E8A105A75FAFFFEB0A06D" lastPageId="5" lastPageNumber="5" pageId="4" pageNumber="4" type="diagnosis">
<paragraph id="1A5736E0FA7E8A115A75FAFFFD0DA024" blockId="4.[573,1012,1349,1430]" box="[573,676,1349,1368]" pageId="4" pageNumber="4">
<heading id="411F818CFA7E8A115A75FAFFFD0DA024" bold="true" box="[573,676,1349,1368]" fontSize="8" level="8" pageId="4" pageNumber="4" reason="0">
<emphasis id="289CEAF2FA7E8A115A75FAFFFD0DA024" bold="true" box="[573,676,1349,1368]" italics="true" pageId="4" pageNumber="4">Diagnosis</emphasis>
</heading>
</paragraph>
<paragraph id="1A5736E0FA7E8A115A75FAD2FC5DA0EA" blockId="4.[573,1012,1349,1430]" pageId="4" pageNumber="4">Small-bodied catarrhine primate (estimated female BM of 4 to 5 kg). Dental formula 2.1.2.3.</paragraph>
<paragraph id="1A5736E0FA7E8A115C50FF3DFB67A64F" blockId="4.[1048,1487,135,1430]" pageId="4" pageNumber="4">Female upper canines moderately compressed. Upper cheek teeth low-crowned and with subpyramidal, moderately peripheral, and inflated cusps. Upper premolars relatively broad and ovoid, P4 smaller than P3, both with heteromorphic cusps, a markedly convex lingual contour and a distinct lingual cingulum (more developed in the P4), a distinct transverse crest separating the restricted mesial fovea from the extensive trigon basin, and the postparacrista forming an abrupt angle with the distal marginal ridge. Upper molars only moderately broader than long, with markedly convex lingual profiles; buccal cusps quite peripheral and buccal cingula discontinuous; lingual cingula relatively well developed, shelf-like, and C-shaped, but not surrounding the hypocone (which is distinct and more peripheral than the protocone); mesial fovea restricted, with an obliquely directed preprotocrista, and trigon basin extensive, being separated by a continuous crista obliqua from the slightly smaller distal fovea, which displays no hypocone-metacone crest. M2 slightly larger than the M1, and M3 shorter and trapezoidal (due to the oblique buccal margin, with a centrally situated metacone and a rudimentary hypocone).</paragraph>
<paragraph id="1A5736E0FA7E8A115C65FC80FA9FA057" blockId="4.[1048,1487,135,1430]" pageId="4" pageNumber="4">Face small but with a distinct snout, the anterior portion of the nasals being almost parallel to the palate. Maxillary sinus large and frontal sinus present but small. Nasal aperture narrow. Nasoalveolar clivus short, with an open palatine fenestra. Anteriorly slightly narrow palate with somewhat convergent upper tooth rows. Zygomatic root moderately high. Orbits subcircular, large, and frontated, with telescopic orbital rims located over the P4. Estimated cranial capacity (69 to 75 cm3) indicating a monkey-like degree of encephalization. External auditory meatus tubular but short and not completely ossified, with a V-shaped end and its anterior portion fused with the postglenoid process. Carotid foramen perforating the bulla posterodistally, and carotid canal horizontally and anteriorly oriented. Spinosum and postglenoid foramina absent. Jugular foramen large and ventrally visible.</paragraph>
<paragraph id="1A5736E0FA7E8A105C65FA89FF62A73F" blockId="4.[1048,1487,135,1430]" lastBlockId="5.[98,538,135,579]" lastPageId="5" lastPageNumber="5" pageId="4" pageNumber="4">Humerus without entepicondylar foramen and capitular tail, with a well-developed capitulum, and a narrow and deep zona conoidea.Radial head rounded and not very tilted, with a markedly beveled surface for articulation with the humeral zona conoidea, the articular surface for the ulnar radial notch extending along a large portion of the radial head, and a laterally facing bicipital tuberosity. Distal radioulnar joint fully diarthrodial, with an expanded and two-faceted semilunar articulation on the ulnar head, and a partially developed ulnar fovea. Ulnar styloid process with reduced girth and not articulating with the short pisiform. Triquetrum small and with a reduced articular surface for the ulnar styloid process. Hamate relatively long proximodistally, with a steep triquetrum facet, a relatively large head and a distally projecting hamulus. Capitate with a relatively small and oblong head and a divided facet for the second metacarpal on its radial side.</paragraph>
<paragraph id="1A5736E0FA7F8A10582AFDE7FEEEA70C" blockId="5.[98,536,605,1297]" box="[98,327,605,624]" pageId="5" pageNumber="5">
<heading id="411F818CFA7F8A10582AFDE7FEEEA70C" bold="true" box="[98,327,605,624]" fontSize="8" level="8" pageId="5" pageNumber="5" reason="0">
<emphasis id="289CEAF2FA7F8A10582AFDE7FEEEA70C" bold="true" box="[98,327,605,624]" italics="true" pageId="5" pageNumber="5">Differential diagnosis</emphasis>
</heading>
</paragraph>
<paragraph id="1A5736E0FA7F8A10582AFD3AFEB0A06D" blockId="5.[98,536,605,1297]" pageId="5" pageNumber="5">
The new taxon differs from pliopithecoids and dendropithecids in its lack of a humeral capitular tail, its hominoid-like proximal radial morphology, its expanded ulnar head with a two-faceted semilunar articulation, and its partially developed ulnar fovea. It further differs from these taxa and proconsulids in its more hominoid-like carpal morphology (including the lack of a pisiform facet for the styloid process, a capitate facet for the second metacarpal divided by a deep ligamentary notch, and a distally projecting hamulus in the hamate), and particularly from pliopithecoids in its overall larger muzzle, more horizontal nasals anteriorly, some details of the upper molars, and (at least compared with
<taxonomicName id="DDE84D63FA7F8A105973FC4EFE6DA17B" box="[315,452,1012,1031]" pageId="5" pageNumber="5">
<emphasis id="289CEAF2FA7F8A105973FC4EFE6DA17B" bold="true" box="[315,452,1012,1031]" italics="true" pageId="5" pageNumber="5">Epipliopithecus</emphasis>
</taxonomicName>
) the lack of an entepicondylar foramen in the humerus. It also differs from all of the above-mentioned taxa in its fused ectotympanic and postglenoid process, and from these taxa and hominids in its horizontal and anteriorly oriented carotid canal. Last, it differs from crown hominoids (hylobatids and hominids) in its incompletely ossified ectotympanic and in its more primitive dentition and forelimb morphology (particularly in the humeroulnar articulation).
</paragraph>
</subSubSection>
<subSubSection id="52F2656BFA7F8A1D582AFA90FC1AA479" lastPageId="8" lastPageNumber="8" pageId="5" pageNumber="5" type="description">
<paragraph id="1A5736E0FA7F8A10582AFA90FE23A024" blockId="5.[98,537,1322,1908]" pageId="5" pageNumber="5">
<heading id="411F818CFA7F8A10582AFA90FE23A024" bold="true" centered="true" fontSize="8" level="7" pageId="5" pageNumber="5" reason="0">
<emphasis id="289CEAF2FA7F8A10582AFA90FE23A024" bold="true" pageId="5" pageNumber="5">
Description, comparisons, and paleobiology
<emphasis id="289CEAF2FA7F8A10582AFAFFFE23A024" bold="true" box="[98,394,1349,1368]" italics="true" pageId="5" pageNumber="5">Dental morphology and diet</emphasis>
</emphasis>
</heading>
</paragraph>
<paragraph id="1A5736E0FA7F8A10582AFAD2FD50A755" blockId="5.[98,537,1322,1908]" lastBlockId="5.[573,1011,135,712]" pageId="5" pageNumber="5">
Although the lack of lower dentition precludes comparisons with some taxa, the upper cheek teeth of
<taxonomicName id="DDE84D63FA7F8A1058E6FA27FF56A0CC" box="[174,255,1437,1456]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7F8A1058E6FA27FF56A0CC" bold="true" box="[174,255,1437,1456]" italics="true" pageId="5" pageNumber="5">Pliobates</emphasis>
</taxonomicName>
(
<figureCitation id="82D32A65FA7F8A105943FA27FE90A0CC" box="[267,313,1437,1456]" captionStart="Fig. 1" captionStartId="2.[98,129,1811,1828]" captionTargetBox="[115,993,753,1778]" captionTargetId="figure@2.[115,993,754,1780]" captionTargetPageId="2" captionText="Fig. 1. Cranium and dentition. (A to C) Cranium of the holotype (IPS 58443) of Pliobates cataloniae gen. et sp. nov. The main cranial fragments, including the basicranium and the right palate, are shown in basal view (A); details of the right palatal fragment are shown in left-lateral (B) and right-lateral (C) views. (D) Detail of the right postcanine teeth, in occlusal view (mesial is to the right)." httpUri="https://zenodo.org/record/269425/files/figure.png" pageId="5" pageNumber="5">Fig. 1</figureCitation>
D and
<tableCitation id="576A035BFA7F8A105930FA27FE11A0CC" box="[376,440,1437,1456]" captionStart="Table 1" captionStartId="4.[605,653,1497,1513]" captionText="Table 1. Dental measurements. Standard dental measurements to assess dental size and proportions were taken to the nearest 0.1 mm in the holotype (IPS 58443) of Pliobates cataloniae gen. et sp. nov. MD, mesiodistal length (in millimeters); BL, buccolingual width (in millimeters); BLI, breadth / length index (in percent), computed as BL / MD × 100. Dashes indicate lack of data due to incomplete preservation." pageId="5" pageNumber="5">Table 1</tableCitation>
) generally resemble those of other small-bodied Miocene catarrhines in both occlusal morphology and proportions (figs. S1 and S2). In contrast, they display a more primitive morphology than those of extant hominoids, including the similarly sized gibbons. Hylobatids possess more elongated cheek teeth with more peripheralized cusps, less developed cingula, and a much more extensive central fovea. Compared with Miocene small-bodied catarrhines from Eurasia and Africa (fig. S1), the upper molars of
<taxonomicName id="DDE84D63FA7F8A105834F97BFF65A3A8" box="[124,204,1729,1748]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7F8A105834F97BFF65A3A8" bold="true" box="[124,204,1729,1748]" italics="true" pageId="5" pageNumber="5">Pliobates</emphasis>
</taxonomicName>
more closely resemble those of the dendropithecid
<emphasis id="289CEAF2FA7F8A1058BCF966FEDDA393" bold="true" box="[244,372,1756,1775]" italics="true" pageId="5" pageNumber="5">Micropithecus</emphasis>
(
<emphasis id="289CEAF2FA7F8A1059CAF966FE3CA393" bold="true" box="[386,405,1756,1775]" italics="true" pageId="5" pageNumber="5">21</emphasis>
,
<emphasis id="289CEAF2FA7F8A1059E9F966FE1EA393" bold="true" box="[417,439,1756,1775]" italics="true" pageId="5" pageNumber="5">34</emphasis>
,
<emphasis id="289CEAF2FA7F8A10598BF966FE71A393" bold="true" box="[451,472,1756,1775]" italics="true" pageId="5" pageNumber="5">35</emphasis>
,
<emphasis id="289CEAF2FA7F8A1059ABF966FE5EA393" bold="true" box="[483,503,1756,1775]" italics="true" pageId="5" pageNumber="5">79</emphasis>
) in several features, such as the markedly convex lingual profiles and moderately developed buccal cingula (albeit to a lesser extent than in
<emphasis id="289CEAF2FA7F8A105992F896FF19A225" bold="true" italics="true" pageId="5" pageNumber="5">Micropithecus</emphasis>
), the C-shaped lingual cingulum that is mostly restricted to the protocone (not surrounding the hypocone), the well-developed and lingually situated hypocone, and the relatively narrow M1 and M2. Nevertheless,
<taxonomicName id="DDE84D63FA7F8A105B42FF07FCF0A5AC" box="[778,857,189,208]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7F8A105B42FF07FCF0A5AC" bold="true" box="[778,857,189,208]" italics="true" pageId="5" pageNumber="5">Pliobates</emphasis>
</taxonomicName>
differs in several features from
<emphasis id="289CEAF2FA7F8A105AF3FF6DFC91A596" bold="true" box="[699,824,215,234]" italics="true" pageId="5" pageNumber="5">Micropithecus</emphasis>
, which has more restricted buccal cingula, a hypocone-metacone crest, and a relatively longer and less trapezoidal M3. The dentition of
<taxonomicName id="DDE84D63FA7F8A105B4CFE9DFCFCA446" box="[772,853,295,314]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7F8A105B4CFE9DFCFCA446" bold="true" box="[772,853,295,314]" italics="true" pageId="5" pageNumber="5">Pliobates</emphasis>
</taxonomicName>
more clearly differs from
<taxonomicName id="DDE84D63FA7F8A105ADAFEFBFCB1A428" box="[658,792,321,340]" pageId="5" pageNumber="5">
<emphasis id="289CEAF2FA7F8A105ADAFEFBFCB1A428" bold="true" box="[658,792,321,340]" italics="true" pageId="5" pageNumber="5">Epipliopithecus</emphasis>
</taxonomicName>
and other pliopithecoids (fig. S1), including from
<emphasis id="289CEAF2FA7F8A105B6CFEE6FC17A413" bold="true" box="[804,958,348,367]" italics="true" pageId="5" pageNumber="5">Barberapithecus</emphasis>
[also recorded at the Vallès-Penedès Basin (
<emphasis id="289CEAF2FA7F8A105BECFECCFC10A4F5" bold="true" box="[932,953,374,393]" italics="true" pageId="5" pageNumber="5">36</emphasis>
)] and
<emphasis id="289CEAF2FA7F8A105A75FE2BFD01A4D8" bold="true" box="[573,680,401,420]" italics="true" pageId="5" pageNumber="5">Pliopithecus</emphasis>
[previously recorded at ACM (
<emphasis id="289CEAF2FA7F8A105BF0FE2BFC64A4D8" bold="true" box="[952,973,401,420]" italics="true" pageId="5" pageNumber="5">33</emphasis>
)] in several features, such as the more convex lingual profile, the more peripheral buccal cusps, the less developed cingula of the molars, the narrower M1 and M2, and the M3 occlusal morphology and proportions (fig. S2).
</paragraph>
<paragraph id="1A5736E0FA7F8A135A1AFD8AFE36A479" blockId="5.[573,1011,135,712]" lastBlockId="6.[98,536,135,261]" lastPageId="6" lastPageNumber="6" pageId="5" pageNumber="5">
With regard to microwear features, the M1 displays a pitting percentage of 30.0%, a pit breadth of 5.67 μm, and a striation breadth of 1.98 μm. Based on pitting incidence (
<figureCitation id="82D32A65FA7F8A105BCCFD3AFC1FA7EF" box="[900,950,640,659]" captionStart="Fig. 2" captionStartId="5.[573,604,1651,1668]" captionTargetBox="[600,1456,769,1637]" captionTargetId="figure@5.[1013,1046,134,1913]" captionTargetPageId="5" captionText="Fig. 2. Results of the dental microwear analyses. (A) Pitting incidence (%) of Pliobates, the extant comparative sample, and pliopithecoids and extinct hominoids from Europe and Turkey. (B) Bivariate plot of striation breadth versus pitting incidence. (C) Bivariate plot of the first two canonical axes delivered by the canonical variates analysis, based on three distinct, broad dietary groups: folivores, mixed feeders and frugivores, and hard-object feeders. Colored polygons in (B) and ellipses in (C) illustrate the variability of extant dietary categories. Small black symbols denote the comparative sample of extant anthropoids, whereas large black symbols represent the centroids of each dietary category. Different symbols are employed to distinguish the various extinct species; results for Iberian hominoids and pliopithecoids are shown in red and blue, respectively, whereas those from other localities are shown in yellow and green, respectively." httpUri="https://zenodo.org/record/269426/files/figure.png" pageId="5" pageNumber="5">Fig. 2</figureCitation>
, A and B), which is the most useful metric for distinguishing among dietary categories (
<emphasis id="289CEAF2FA7F8A105B7DFD0FFCE3A7B4" bold="true" box="[821,842,693,712]" italics="true" pageId="5" pageNumber="5">70</emphasis>
),
<taxonomicName id="DDE84D63FA7F8A105B16FD0FFC06A7B4" box="[862,943,693,712]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7F8A105B16FD0FFC06A7B4" bold="true" box="[862,943,693,712]" italics="true" pageId="5" pageNumber="5">Pliobates</emphasis>
</taxonomicName>
closely resembles extant frugivores (
<taxonomicName id="DDE84D63FA7F8A105D5EFF3DFA36A5E6" box="[1302,1439,135,154]" class="Mammalia" family="Hominidae" genus="Pan" kingdom="Animalia" order="Primates" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="troglodytes">
<emphasis id="289CEAF2FA7F8A105D5EFF3DFA36A5E6" bold="true" box="[1302,1439,135,154]" italics="true" pageId="5" pageNumber="5">Pan troglodytes</emphasis>
</taxonomicName>
) and eclectic feeders (
<taxonomicName id="DDE84D63FA7F8A105CE3FF18FAF2A5C9" box="[1195,1371,162,181]" class="Mammalia" family="Cercopithecidae" genus="Papio" kingdom="Animalia" order="Primates" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="cynocephalus">
<emphasis id="289CEAF2FA7F8A105CE3FF18FAF2A5C9" bold="true" box="[1195,1371,162,181]" italics="true" pageId="5" pageNumber="5">Papio cynocephalus</emphasis>
</taxonomicName>
) that largely rely on ripe fruit. In contrast, the pitting incidence of
<taxonomicName id="DDE84D63FA7F8A105C67FF6DFBD7A596" box="[1071,1150,215,234]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7F8A105C67FF6DFBD7A596" bold="true" box="[1071,1150,215,234]" italics="true" pageId="5" pageNumber="5">Pliobates</emphasis>
</taxonomicName>
is higher than in extant folivores and much lower than in extant hard-object feeders (including orangutans). Compared with other extinct catarrhines from Western Europe, the pitting percentage of
<taxonomicName id="DDE84D63FA7F8A105CD2FEFBFB44A428" box="[1178,1261,321,340]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7F8A105CD2FEFBFB44A428" bold="true" box="[1178,1261,321,340]" italics="true" pageId="5" pageNumber="5">Pliobates</emphasis>
</taxonomicName>
is somewhat lower than in most pliopithecoids and hominoids, for which some degree of sclerocarpy has been inferred (
<emphasis id="289CEAF2FA7F8A105C57FE2BFB9FA4D8" bold="true" box="[1055,1078,401,420]" italics="true" pageId="5" pageNumber="5">66</emphasis>
,
<emphasis id="289CEAF2FA7F8A105C08FE2BFBFEA4D8" bold="true" box="[1088,1111,401,420]" italics="true" pageId="5" pageNumber="5">68</emphasis>
). This low pitting incidence is consistent with the pit- and striation-breadth measurements (which show no sign of extreme folivory or specialized hard-object feeding) and most closely approaches that of the fossil hominoids
<emphasis id="289CEAF2FA7F8A105C50FDACFAAEA755" bold="true" box="[1048,1287,534,553]" italics="true" pageId="5" pageNumber="5">Anoiapithecus brevirostris</emphasis>
and
<emphasis id="289CEAF2FA7F8A105D7EFDACFBC5A73F" bold="true" italics="true" pageId="5" pageNumber="5">Hispanopithecus laietanus</emphasis>
, previously interpreted as soft frugivores (
<emphasis id="289CEAF2FA7F8A105C57FDF1FB9CA722" bold="true" box="[1055,1077,587,606]" italics="true" pageId="5" pageNumber="5">68</emphasis>
). These results are confirmed by a multivariate analysis that simultaneously examined the three microwear variables (
<figureCitation id="82D32A65FA7F8A105C9DFD3AFAACA7EF" box="[1237,1285,640,659]" captionStart="Fig. 2" captionStartId="5.[573,604,1651,1668]" captionTargetBox="[600,1456,769,1637]" captionTargetId="figure@5.[1013,1046,134,1913]" captionTargetPageId="5" captionText="Fig. 2. Results of the dental microwear analyses. (A) Pitting incidence (%) of Pliobates, the extant comparative sample, and pliopithecoids and extinct hominoids from Europe and Turkey. (B) Bivariate plot of striation breadth versus pitting incidence. (C) Bivariate plot of the first two canonical axes delivered by the canonical variates analysis, based on three distinct, broad dietary groups: folivores, mixed feeders and frugivores, and hard-object feeders. Colored polygons in (B) and ellipses in (C) illustrate the variability of extant dietary categories. Small black symbols denote the comparative sample of extant anthropoids, whereas large black symbols represent the centroids of each dietary category. Different symbols are employed to distinguish the various extinct species; results for Iberian hominoids and pliopithecoids are shown in red and blue, respectively, whereas those from other localities are shown in yellow and green, respectively." httpUri="https://zenodo.org/record/269426/files/figure.png" pageId="5" pageNumber="5">Fig. 2</figureCitation>
C and tables S9 to S11), in which
<taxonomicName id="DDE84D63FA7F8A105C32FD21FB67A7D2" box="[1146,1230,667,686]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7F8A105C32FD21FB67A7D2" bold="true" box="[1146,1230,667,686]" italics="true" pageId="5" pageNumber="5">Pliobates</emphasis>
</taxonomicName>
falls closer to the extant frugivorousmixed-feeder centroid for the first and second canonical axes and is classified as a frugivore. Dental microwear analyses therefore indicate a mainly frugivorous diet for
<taxonomicName id="DDE84D63FA7C8A13598AFF07FDBAA5AC" box="[450,531,189,208]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7C8A13598AFF07FDBAA5AC" bold="true" box="[450,531,189,208]" italics="true" pageId="6" pageNumber="6">Pliobates</emphasis>
</taxonomicName>
, compatible with a high consumption of ripe fruit and a low sclerocarpic component.
</paragraph>
<caption id="4E976668FA7F8A105A75F9C9FCFBA20F" httpUri="https://zenodo.org/record/269426/files/figure.png" pageId="5" pageNumber="5" targetBox="[600,1456,769,1637]" targetPageId="5">
<paragraph id="1A5736E0FA7F8A105A75F9C9FCFBA20F" blockId="5.[573,1487,1650,1907]" pageId="5" pageNumber="5">
<emphasis id="289CEAF2FA7F8A105A75F9C9FBBCA3F8" bold="true" box="[573,1045,1651,1668]" pageId="5" pageNumber="5">Fig. 2. Results of the dental microwear analyses.</emphasis>
(
<emphasis id="289CEAF2FA7F8A105C6BF9C9FB98A3F8" bold="true" box="[1059,1073,1651,1668]" pageId="5" pageNumber="5">A</emphasis>
) Pitting incidence (%) of
<taxonomicName id="DDE84D63FA7F8A105D50F9C9FAC0A3F8" box="[1304,1385,1651,1668]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="5" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7F8A105D50F9C9FAC0A3F8" box="[1304,1385,1651,1668]" italics="true" pageId="5" pageNumber="5">Pliobates</emphasis>
</taxonomicName>
, the extant comparative sample, and pliopithecoids and extinct hominoids from Europe and Turkey. (
<emphasis id="289CEAF2FA7F8A105D2BF934FAD8A3E2" bold="true" box="[1379,1393,1678,1694]" pageId="5" pageNumber="5">B</emphasis>
) Bivariate plot of striation breadth versus pitting incidence. (
<emphasis id="289CEAF2FA7F8A105C5AF912FB89A3C5" bold="true" box="[1042,1056,1704,1721]" pageId="5" pageNumber="5">C</emphasis>
) Bivariate plot of the first two canonical axes delivered by the canonical variates analysis, based on three distinct, broad dietary groups: folivores, mixed feeders and frugivores, and hard-object feeders. Colored polygons in (B) and ellipses in (C) illustrate the variability of extant dietary categories. Small black symbols denote the comparative sample of extant anthropoids, whereas large black symbols represent the centroids of each dietary category. Different symbols are employed to distinguish the various extinct species; results for Iberian hominoids and pliopithecoids are shown in red and blue, respectively,whereas those from other localities are shown in yellow and green, respectively.
</paragraph>
</caption>
<paragraph id="1A5736E0FA7C8A13582AFEA4FF7CA44E" blockId="6.[98,536,286,819]" box="[98,213,286,306]" pageId="6" pageNumber="6">
<heading id="411F818CFA7C8A13582AFEA4FF7CA44E" bold="true" box="[98,213,286,306]" fontSize="8" level="8" pageId="6" pageNumber="6" reason="0">
<emphasis id="289CEAF2FA7C8A13582AFEA4FF7CA44E" bold="true" box="[98,213,286,306]" italics="true" pageId="6" pageNumber="6">Body mass</emphasis>
</heading>
</paragraph>
<paragraph id="1A5736E0FA7C8A13582AFEFBFE51A64F" blockId="6.[98,536,286,819]" pageId="6" pageNumber="6">
Dental BM estimates for the female holotype of
<taxonomicName id="DDE84D63FA7C8A13582AFEE6FF1AA413" box="[98,179,348,367]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7C8A13582AFEE6FF1AA413" bold="true" box="[98,179,348,367]" italics="true" pageId="6" pageNumber="6">Pliobates</emphasis>
</taxonomicName>
(table S7) range from 2.9 to 4.8 kg, with an average BM estimate of 3.9 kg and an uncertainty degree (based on the combined 95% confidence intervals for each dental locus) of 2.5 to 5.7 kg. Postcranial BM estimates (table S8) are on average 4.8 kg (range: 4.0 to 5.6 kg), based on catarrhine regressions, and 4.3 kg (range: 2.6 to 6.3 kg), based on hominoid regressions (estimates for each postcranial estimator and their confidence intervals are given in table S8). Given that the size of
<taxonomicName id="DDE84D63FA7C8A1358F5FDDCFEA3A705" box="[189,266,614,633]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7C8A1358F5FDDCFEA3A705" bold="true" box="[189,266,614,633]" italics="true" pageId="6" pageNumber="6">Pliobates</emphasis>
</taxonomicName>
is in the lower range for extant hominoids, the catarrhine regressions probably yield more accurate estimates, although the hominoid-based estimates are closer to the dental ones. Overall, the body mass of the holotype of
<taxonomicName id="DDE84D63FA7C8A13582AFD50FEA2A781" box="[98,267,746,765]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="cataloniae">
<emphasis id="289CEAF2FA7C8A13582AFD50FEA2A781" bold="true" box="[98,267,746,765]" italics="true" pageId="6" pageNumber="6">Pliobates cataloniae</emphasis>
</taxonomicName>
can be estimated at ~4 to 5 kg (much lower than that estimated for
<taxonomicName id="DDE84D63FA7C8A1359F5FCBFFE8BA64F" authority="Zapfe &amp; Hürzeler, 1957" authorityName="Zapfe &amp; Hürzeler" authorityYear="1957" class="Mammalia" genus="Epipliopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="vindobonensis">
<emphasis id="289CEAF2FA7C8A1359F5FCBFFE8BA64F" bold="true" italics="true" pageId="6" pageNumber="6">Epipliopithecus vindobonensis</emphasis>
</taxonomicName>
, ~11 to 12 kg; table S8).
</paragraph>
<paragraph id="1A5736E0FA7C8A13582AFCF6FDA4A623" blockId="6.[98,537,844,1908]" box="[98,525,844,863]" pageId="6" pageNumber="6">
<heading id="411F818CFA7C8A13582AFCF6FDA4A623" bold="true" box="[98,525,844,863]" fontSize="8" level="8" pageId="6" pageNumber="6" reason="0">
<emphasis id="289CEAF2FA7C8A13582AFCF6FDA4A623" bold="true" box="[98,525,844,863]" italics="true" pageId="6" pageNumber="6">Cranial morphology and encephalization</emphasis>
</heading>
</paragraph>
<paragraph id="1A5736E0FA7C8A13582AFCD5FB18A5AC" blockId="6.[98,537,844,1908]" lastBlockId="6.[573,1485,135,208]" pageId="6" pageNumber="6">
A 3D virtual reconstruction of the cranium, based on the preserved specimens, is shown in fig. S3, whereas the final reconstruction (including mirrored portions) is shown in
<figureCitation id="82D32A65FA7C8A13592AFC05FE3DA6AE" box="[354,404,959,978]" captionStart="Fig. 3" captionStartId="6.[573,603,907,924]" captionTargetBox="[591,1468,257,876]" captionTargetId="figure@6.[591,1469,257,876]" captionTargetPageId="6" captionText="Fig. 3. Cranial reconstruction. Virtual reconstruction of the holotype (IPS 58443) cranium of Pliobates cataloniae gen. et sp. nov., including mirrored fragments, in frontal (A), lateral (B), posterior (C), basal (D), and superior (E) views. Further details are given in fig. S 3 and the methods in the text." httpUri="https://zenodo.org/record/269427/files/figure.png" pageId="6" pageNumber="6">Fig. 3</figureCitation>
and movie S1. Based on this reconstruction, the cranium of
<taxonomicName id="DDE84D63FA7C8A13582AFC4EFF19A17B" box="[98,176,1012,1031]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7C8A13582AFC4EFF19A17B" bold="true" box="[98,176,1012,1031]" italics="true" pageId="6" pageNumber="6">Pliobates</emphasis>
</taxonomicName>
differs from the primitive catarrhine condition (
<emphasis id="289CEAF2FA7C8A1358ECFBB5FF10A15E" bold="true" box="[164,185,1039,1058]" italics="true" pageId="6" pageNumber="6">22</emphasis>
,
<emphasis id="289CEAF2FA7C8A13588CFBB5FF71A15E" bold="true" box="[196,216,1039,1058]" italics="true" pageId="6" pageNumber="6">37</emphasis>
,
<emphasis id="289CEAF2FA7C8A1358A9FBB5FF5EA15E" bold="true" box="[225,247,1039,1058]" italics="true" pageId="6" pageNumber="6">60</emphasis>
) by being short, wide, and high. However, the tubular ectotympanic is short and incompletely ossified—i.e., less developed than in
<emphasis id="289CEAF2FA7C8A13582AFBE4FF68A10D" bold="true" box="[98,193,1118,1137]" italics="true" pageId="6" pageNumber="6">Saadanius</emphasis>
and extant crown catarrhines (
<emphasis id="289CEAF2FA7C8A13599FFBE4FDA5A10D" bold="true" box="[471,524,1118,1137]" italics="true" pageId="6" pageNumber="6">2022</emphasis>
). The maxillary sinus is extensive, as in stem catarrhines and hominoids (
<emphasis id="289CEAF2FA7C8A13591DFB2EFEC3A1DB" bold="true" box="[341,362,1172,1191]" italics="true" pageId="6" pageNumber="6">22</emphasis>
,
<emphasis id="289CEAF2FA7C8A135930FB2EFE26A1DB" bold="true" box="[376,399,1172,1191]" italics="true" pageId="6" pageNumber="6">80</emphasis>
), and there is also a small frontal sinus, as in stem hominoids but unlike in stem catarrhines, cercopithecoids, hylobatids, and pongines (
<emphasis id="289CEAF2FA7C8A135919FB59FECFA18A" bold="true" box="[337,358,1251,1270]" italics="true" pageId="6" pageNumber="6">22</emphasis>
,
<emphasis id="289CEAF2FA7C8A135939FB59FE2CA18A" bold="true" box="[369,389,1251,1270]" italics="true" pageId="6" pageNumber="6">37</emphasis>
,
<emphasis id="289CEAF2FA7C8A1359C7FB59FE0FA18A" bold="true" box="[399,422,1251,1270]" italics="true" pageId="6" pageNumber="6">80</emphasis>
). The face is short and displays anteriorly situated orbits, as in hylobatids, colobines, and some extinct small-bodied catarrhines such as
<taxonomicName id="DDE84D63FA7C8A135947FA89FE38A03A" box="[271,401,1331,1350]" pageId="6" pageNumber="6">
<emphasis id="289CEAF2FA7C8A135947FA89FE38A03A" bold="true" box="[271,401,1331,1350]" italics="true" pageId="6" pageNumber="6">Epipliopithecus</emphasis>
</taxonomicName>
,
<emphasis id="289CEAF2FA7C8A1359D4FA89FDBAA03A" bold="true" box="[412,531,1331,1350]" italics="true" pageId="6" pageNumber="6">Micropithecus</emphasis>
, and
<emphasis id="289CEAF2FA7C8A1358C2FAF7FEB4A01C" bold="true" box="[138,285,1357,1376]" italics="true" pageId="6" pageNumber="6">Lomorupithecus</emphasis>
(
<emphasis id="289CEAF2FA7C8A135960FAF7FE92A01C" bold="true" box="[296,315,1357,1376]" italics="true" pageId="6" pageNumber="6">19</emphasis>
,
<emphasis id="289CEAF2FA7C8A13590DFAF7FEF1A01C" bold="true" box="[325,344,1357,1376]" italics="true" pageId="6" pageNumber="6">21</emphasis>
,
<emphasis id="289CEAF2FA7C8A135929FAF7FEDEA01C" bold="true" box="[353,375,1357,1376]" italics="true" pageId="6" pageNumber="6">38</emphasis>
,
<emphasis id="289CEAF2FA7C8A1359C9FAF7FE3CA01C" bold="true" box="[385,405,1357,1376]" italics="true" pageId="6" pageNumber="6">76</emphasis>
,
<emphasis id="289CEAF2FA7C8A1359D6FAF7FE1BA01C" bold="true" box="[414,434,1357,1376]" italics="true" pageId="6" pageNumber="6">79</emphasis>
). However,
<taxonomicName id="DDE84D63FA7C8A13582AFAD2FF1AA007" box="[98,179,1384,1403]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7C8A13582AFAD2FF1AA007" bold="true" box="[98,179,1384,1403]" italics="true" pageId="6" pageNumber="6">Pliobates</emphasis>
</taxonomicName>
differs from these taxa (and more closely resembles hylobatids) by displaying a more well-defined muzzle (especially compared with
<taxonomicName id="DDE84D63FA7C8A13582AFA02FF46A0B7" box="[98,239,1464,1483]" pageId="6" pageNumber="6">
<emphasis id="289CEAF2FA7C8A13582AFA02FF46A0B7" bold="true" box="[98,239,1464,1483]" italics="true" pageId="6" pageNumber="6">Epipliopithecus</emphasis>
</taxonomicName>
) with long and more horizontal nasals, a higher zygomatic root (moderately high as in hylobatids, but less so than in hominids), an interorbital pillar nearly orthogonal to the frontal squama (as in hylobatids and chimpanzees), a high degree of orbital convergence and frontation (as in all extant hominoids), and thin and anteriorly projecting (telescopic) orbital rims [to a greater extent than in
<taxonomicName id="DDE84D63FA7C8A13597FF936FE6DA3E3" box="[311,452,1676,1695]" pageId="6" pageNumber="6">
<emphasis id="289CEAF2FA7C8A13597FF936FE6DA3E3" bold="true" box="[311,452,1676,1695]" italics="true" pageId="6" pageNumber="6">Epipliopithecus</emphasis>
</taxonomicName>
(
<emphasis id="289CEAF2FA7C8A135987F936FE4CA3E3" bold="true" box="[463,485,1676,1695]" italics="true" pageId="6" pageNumber="6">38</emphasis>
), and thus most closely resembling hylobatids and, as far as it can be ascertained with incomplete preservation,
<emphasis id="289CEAF2FA7C8A135888F966FEE9A393" bold="true" box="[192,320,1756,1775]" italics="true" pageId="6" pageNumber="6">Micropithecus</emphasis>
(
<emphasis id="289CEAF2FA7C8A135904F966FEC9A393" bold="true" box="[332,352,1756,1775]" italics="true" pageId="6" pageNumber="6">79</emphasis>
)].
<taxonomicName id="DDE84D63FA7C8A135930F966FE60A393" box="[376,457,1756,1775]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7C8A135930F966FE60A393" bold="true" box="[376,457,1756,1775]" italics="true" pageId="6" pageNumber="6">Pliobates</emphasis>
</taxonomicName>
also displays derived hominoid features in the basicranium (
<figureCitation id="82D32A65FA7C8A135821F8ABFF32A258" box="[105,155,1809,1828]" captionStart="Fig. 4" captionStartId="6.[573,604,1625,1642]" captionTargetBox="[592,1468,1026,1624]" captionTargetId="figure@6.[590,1468,1026,1610]" captionTargetPageId="6" captionText="Fig. 4. Basicranial morphology. (A) Drawing of the left basicranium of the holotype (IPS 58443) of Pliobates cataloniae gen. et sp. nov., as preserved in ventral view. The jugular foramen appears artifactually larger because of the displacement of the temporal and occipital portions along the occipitotemporal suture (corrected in the reconstruction in Fig. 3). The course of the carotid canal is shown with a dashed line, based on CT images. AE, articular eminence; CAF, carotid foramen; COF, condylar fossa; EA, Eustachian aperture; EAM, external auditory meatus; EP, Eustachian process; ET, ectotympanic; FM, foramen magnum; FO, foramen ovale; GF, glenoid fossa; JF, jugular foramen; OC, occipital condyle; OTS, occipitotemporal suture; PGP, postglenoid process. (B to D) Drawings of comparable views (not to scale) in Hylobates sp. (B), Proconsul heseloni KNM RU 2036 [(C), reversed], and Victoriapithecus macinnesi KNM MB 29100 a (D) (KNM, Kenyon National Museums; RU, Rusinga; MB, Maboko). Arrows denote the V-shaped, incompletely ossified ventral terminal tip of the tubular ectotympanic in the extinct taxa. [Artwork by M. Palmero]" httpUri="https://zenodo.org/record/269428/files/figure.png" pageId="6" pageNumber="6">Fig. 4</figureCitation>
A), including the absence of a postglenoid foramen with a large and ventrally visible jugular foramen (as in all extant hominoids), the foramen ovale situated anteriorly and laterally to the Eusta- chian aperture (as in hylobatids and African and African apes), and the horizontal and anapes), the fusion between the auditory meatus teriorly directed carotid canal in the petrosal bone and the postglenoid process (as in hylobatids (as in hylobatids).
</paragraph>
<caption id="4E976668FA7C8A135A75FC31FAC0A6AD" httpUri="https://zenodo.org/record/269427/files/figure.png" pageId="6" pageNumber="6" targetBox="[591,1468,257,876]" targetPageId="6">
<paragraph id="1A5736E0FA7C8A135A75FC31FAC0A6AD" blockId="6.[573,1485,906,978]" pageId="6" pageNumber="6">
<emphasis id="289CEAF2FA7C8A135A75FC31FCFBA6E0" bold="true" box="[573,850,907,924]" pageId="6" pageNumber="6">Fig. 3. Cranial reconstruction.</emphasis>
Virtual reconstruction of the holotype (IPS58443) cranium of
<taxonomicName id="DDE84D63FA7C8A135D36FC31FD3FA6CB" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="cataloniae">
<emphasis id="289CEAF2FA7C8A135D36FC31FD3FA6CB" italics="true" pageId="6" pageNumber="6">Pliobates cataloniae</emphasis>
</taxonomicName>
gen. et sp. nov., including mirrored fragments, in frontal (
<emphasis id="289CEAF2FA7C8A135CD2FC1CFB01A6CB" bold="true" box="[1178,1192,934,951]" pageId="6" pageNumber="6">A</emphasis>
), lateral (
<emphasis id="289CEAF2FA7C8A135D48FC1CFAA7A6CA" bold="true" box="[1280,1294,934,950]" pageId="6" pageNumber="6">B</emphasis>
), posterior (
<emphasis id="289CEAF2FA7C8A135D36FC1CFA25A6CB" bold="true" box="[1406,1420,934,951]" pageId="6" pageNumber="6">C</emphasis>
), basal (
<emphasis id="289CEAF2FA7C8A135A0DFC7BFDFAA6AD" bold="true" box="[581,595,961,977]" pageId="6" pageNumber="6">D</emphasis>
), and superior (
<emphasis id="289CEAF2FA7C8A135AABFC7BFD46A6AD" bold="true" box="[739,751,961,977]" pageId="6" pageNumber="6">E</emphasis>
) views. Further details are given in fig. S3 and the methods in the text.
</paragraph>
</caption>
<caption id="4E976668FA7C8A135A75F9E3FA02A208" httpUri="https://zenodo.org/record/269428/files/figure.png" pageId="6" pageNumber="6" targetBox="[592,1468,1026,1624]" targetPageId="6">
<paragraph id="1A5736E0FA7C8A135A75F9E3FA02A208" blockId="6.[573,1486,1623,1908]" pageId="6" pageNumber="6">
<emphasis id="289CEAF2FA7C8A135A75F9E3FCC0A315" bold="true" box="[573,873,1625,1642]" pageId="6" pageNumber="6">Fig. 4. Basicranial morphology.</emphasis>
(
<emphasis id="289CEAF2FA7C8A135B3EF9E3FC2DA316" bold="true" box="[886,900,1625,1642]" pageId="6" pageNumber="6">A</emphasis>
) Drawing of the left basicranium of the holotype (IPS58443) of
<taxonomicName id="DDE84D63FA7C8A135A75F9C9FD4EA3F8" box="[573,743,1651,1668]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="cataloniae">
<emphasis id="289CEAF2FA7C8A135A75F9C9FD4EA3F8" box="[573,743,1651,1668]" italics="true" pageId="6" pageNumber="6">Pliobates cataloniae</emphasis>
</taxonomicName>
gen. et sp. nov., as preserved in ventral view.The jugular foramen appears artifactually larger because of the displacement of the temporal and occipital portions along the occipitotemporal suture (corrected in the reconstruction in Fig. 3).The course of the carotid canal is shown with a dashed line, based on CT images. AE, articular eminence; CAF, carotid foramen; COF, condylar fossa; EA, Eustachian aperture; EAM, external auditory meatus; EP, Eustachian process; ET, ectotympanic; FM, foramen magnum; FO, foramen ovale; GF, glenoid fossa; JF, jugular foramen; OC, occipital condyle; OTS, occipitotemporal suture; PGP, postglenoid process. (
<emphasis id="289CEAF2FA7C8A135B25F8A9FCD2A25F" bold="true" box="[877,891,1811,1827]" pageId="6" pageNumber="6">B</emphasis>
to
<emphasis id="289CEAF2FA7C8A135BD3F8A9FC00A25F" bold="true" box="[923,937,1811,1827]" pageId="6" pageNumber="6">D</emphasis>
) Drawings of comparable views (not to scale) in
<taxonomicName id="DDE84D63FA7C8A135D12F8A9FA07A258" box="[1370,1454,1811,1828]" class="Mammalia" family="Hylobatidae" genus="Hylobates" kingdom="Animalia" order="Primates" pageId="6" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="289CEAF2FA7C8A135D12F8A9FA07A258" box="[1370,1454,1811,1828]" italics="true" pageId="6" pageNumber="6">Hylobates</emphasis>
</taxonomicName>
sp. (B),
<emphasis id="289CEAF2FA7C8A135A2DF897FCACA242" box="[613,773,1837,1854]" italics="true" pageId="6" pageNumber="6">Proconsul heseloni</emphasis>
KNM RU 2036 [(C), reversed], and
<emphasis id="289CEAF2FA7C8A135C01F897FA85A242" box="[1097,1324,1837,1854]" italics="true" pageId="6" pageNumber="6">Victoriapithecus macinnesi</emphasis>
KNM MB 29100a (D) (KNM, Kenyon National Museums; RU, Rusinga; MB, Maboko). Arrows denote the V-shaped, incompletely ossified ventral terminal tip of the tubular ectotympanic in the extinct taxa. [Artwork by M. Palmero]
</paragraph>
</caption>
<paragraph id="1A5736E0FA7D8A125830FF3DFDB1A664" blockId="7.[98,536,135,792]" pageId="7" pageNumber="7">
Braincase measurements yield an average cranial capacity estimate of 69.0 cm3 (range: 41.4 to 110.7 cm 3; table S12), which is close to the estimates of 60.1 and 65.3 cm3 delivered by the two most reliable estimators (
<emphasis id="289CEAF2FA7D8A12591DFF48FEC1A479" bold="true" box="[341,360,242,261]" italics="true" pageId="7" pageNumber="7">57</emphasis>
) and only slightly lower than the estimate of 75.1 cm3 obtained from foramen magnum area (table S12). According to our estimates of body mass (4.5 kg) and cranial capacity [72 cm3 (average of cranial and foramen magnum estimates)],
<taxonomicName id="DDE84D63FA7D8A125964FECCFED4A4F5" box="[300,381,374,393]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="7" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7D8A125964FECCFED4A4F5" bold="true" box="[300,381,374,393]" italics="true" pageId="7" pageNumber="7">Pliobates</emphasis>
</taxonomicName>
would display a monkey-like degree of encephalization extensively overlapping with extant cercopithecoids (fig. S4 and table S13), being much more encephalized than the stem catarrhine
<emphasis id="289CEAF2FA7D8A125958FE5BFE37A488" bold="true" box="[272,414,481,500]" italics="true" pageId="7" pageNumber="7">Aegyptopithecus</emphasis>
, slightly more so than the stem cercopithecoid
<emphasis id="289CEAF2FA7D8A1259CDFE41FDBAA772" bold="true" box="[389,531,507,526]" italics="true" pageId="7" pageNumber="7">Victoriapithecus</emphasis>
, and only slightly less so than hylobatids and the extinct hominoids
<emphasis id="289CEAF2FA7D8A12594CFD8AFEF3A73F" bold="true" box="[260,346,560,579]" italics="true" pageId="7" pageNumber="7">Proconsul</emphasis>
and
<emphasis id="289CEAF2FA7D8A1259CDFD8AFE5FA73F" bold="true" box="[389,502,560,579]" italics="true" pageId="7" pageNumber="7">Oreopithecus</emphasis>
. All these taxa, like cercopithecoids, are less encephalized than the extinct hominoid
<emphasis id="289CEAF2FA7D8A1259CBFDDCFDB1A705" bold="true" box="[387,536,614,633]" italics="true" pageId="7" pageNumber="7">Hispanopithecus</emphasis>
(
<emphasis id="289CEAF2FA7D8A125821FD3AFF77A7EF" bold="true" box="[105,222,640,659]" italics="true" pageId="7" pageNumber="7">Rudapithecus</emphasis>
) and the extant great apes. Although humans are outliers in brain sizebody size allometric regressions, great apes further display an allometric grade shift compared with hylobatids (and
<taxonomicName id="DDE84D63FA7D8A1258D9FD50FF4BA781" box="[145,226,746,765]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="7" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7D8A1258D9FD50FF4BA781" bold="true" box="[145,226,746,765]" italics="true" pageId="7" pageNumber="7">Pliobates</emphasis>
</taxonomicName>
), which are only slightly more encephalized on average than cercopithecoids (
<emphasis id="289CEAF2FA7D8A1259BFFCBFFDA5A664" bold="true" box="[503,524,773,792]" italics="true" pageId="7" pageNumber="7">58</emphasis>
).
</paragraph>
<paragraph id="1A5736E0FA7D8A12582AFC8BFDA3A638" blockId="7.[98,537,817,1908]" box="[98,522,817,836]" pageId="7" pageNumber="7">
<heading id="411F818CFA7D8A12582AFC8BFDA3A638" bold="true" box="[98,522,817,836]" fontSize="8" level="8" pageId="7" pageNumber="7" reason="0">
<emphasis id="289CEAF2FA7D8A12582AFC8BFDA3A638" bold="true" box="[98,522,817,836]" italics="true" pageId="7" pageNumber="7">Postcranial morphology and locomotion</emphasis>
</heading>
</paragraph>
<paragraph id="1A5736E0FA7D8A12582AFCEFFE24A393" blockId="7.[98,537,817,1908]" pageId="7" pageNumber="7">
The humerus (
<figureCitation id="82D32A65FA7D8A125895FCEFFEA2A614" box="[221,267,853,872]" captionStart="Fig. 5" captionStartId="7.[573,604,1758,1775]" captionTargetBox="[590,1468,989,1726]" captionTargetId="figure@7.[1013,1046,134,1913]" captionTargetPageId="7" captionText="Fig. 5. Forelimb long bones. Shown are the humerus, radius, and ulna of the holotype (IPS 58443) of Pliobates cataloniae gen. et sp. nov. (A to E) Partial left humerus in medial (A), posterior (B), lateral (C), anterior (D), and distal (E) views. (F to K) Left radius in medial (F), posterior (G), lateral (H), anterior (I), proximal (J), and distal (K) views. (L to O) Proximal half of the left ulna in medial (L), posterior (M), lateral (N), and anterior (O) views. (P to T) Distal fragment of the left ulna in medial (P), posterior (Q), lateral (R), anterior (S), and distal (T) views." httpUri="https://zenodo.org/record/269429/files/figure.png" pageId="7" pageNumber="7">Fig. 5</figureCitation>
) resembles that of extant crown catarrhines, proconsulids, and dendropithecids by lacking (unlike
<taxonomicName id="DDE84D63FA7D8A1258A0FC30FED9A6E1" authority="Zapfe &amp; Hürzeler, 1957" authorityName="Zapfe &amp; Hürzeler" authorityYear="1957" box="[232,368,906,925]" class="Mammalia" genus="Epipliopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="7" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="289CEAF2FA7D8A1258A0FC30FED9A6E1" bold="true" box="[232,368,906,925]" italics="true" pageId="7" pageNumber="7">Epipliopithecus</emphasis>
</taxonomicName>
) an entepicondylar foramen (
<emphasis id="289CEAF2FA7D8A1258F1FC1EFF60A6CB" bold="true" box="[185,201,932,951]" italics="true" pageId="7" pageNumber="7">17</emphasis>
,
<emphasis id="289CEAF2FA7D8A12589DFC1EFF42A6CB" bold="true" box="[213,235,932,951]" italics="true" pageId="7" pageNumber="7">20</emphasis>
,
<emphasis id="289CEAF2FA7D8A1258BDFC1EFEAEA6CB" bold="true" box="[245,263,932,951]" italics="true" pageId="7" pageNumber="7">21</emphasis>
,
<emphasis id="289CEAF2FA7D8A12595AFC1EFE8EA6CB" bold="true" box="[274,295,932,951]" italics="true" pageId="7" pageNumber="7">38</emphasis>
,
<emphasis id="289CEAF2FA7D8A12597AFC1EFEECA6CB" bold="true" box="[306,325,932,951]" italics="true" pageId="7" pageNumber="7">79</emphasis>
,
<emphasis id="289CEAF2FA7D8A125919FC1EFECDA6CB" bold="true" box="[337,356,932,951]" italics="true" pageId="7" pageNumber="7">81</emphasis>
,
<emphasis id="289CEAF2FA7D8A125926FC1EFE2AA6CB" bold="true" box="[366,387,932,951]" italics="true" pageId="7" pageNumber="7">82</emphasis>
).
<taxonomicName id="DDE84D63FA7D8A1259DDFC1EFE4AA6CB" box="[405,483,932,951]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="7" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7D8A1259DDFC1EFE4AA6CB" bold="true" box="[405,483,932,951]" italics="true" pageId="7" pageNumber="7">Pliobates</emphasis>
</taxonomicName>
more closely resembles extant hominoids in the laterally facing bicipital tuberosity in the radius (
<emphasis id="289CEAF2FA7D8A125993FC60FE47A691" bold="true" box="[475,494,986,1005]" italics="true" pageId="7" pageNumber="7">81</emphasis>
,
<emphasis id="289CEAF2FA7D8A1259B3FC60FDB8A691" bold="true" box="[507,529,986,1005]" italics="true" pageId="7" pageNumber="7">83</emphasis>
) (
<figureCitation id="82D32A65FA7D8A125820FC4EFF36A17B" box="[104,159,1012,1031]" captionStart="Fig. 5" captionStartId="7.[573,604,1758,1775]" captionTargetBox="[590,1468,989,1726]" captionTargetId="figure@7.[1013,1046,134,1913]" captionTargetPageId="7" captionText="Fig. 5. Forelimb long bones. Shown are the humerus, radius, and ulna of the holotype (IPS 58443) of Pliobates cataloniae gen. et sp. nov. (A to E) Partial left humerus in medial (A), posterior (B), lateral (C), anterior (D), and distal (E) views. (F to K) Left radius in medial (F), posterior (G), lateral (H), anterior (I), proximal (J), and distal (K) views. (L to O) Proximal half of the left ulna in medial (L), posterior (M), lateral (N), and anterior (O) views. (P to T) Distal fragment of the left ulna in medial (P), posterior (Q), lateral (R), anterior (S), and distal (T) views." httpUri="https://zenodo.org/record/269429/files/figure.png" pageId="7" pageNumber="7">Fig. 5</figureCitation>
), as well as in the configuration of the humeroradial articulation (
<emphasis id="289CEAF2FA7D8A12592AFBB5FEDCA15E" bold="true" box="[354,373,1039,1058]" italics="true" pageId="7" pageNumber="7">81</emphasis>
,
<emphasis id="289CEAF2FA7D8A1259CBFBB5FE30A15E" bold="true" box="[387,409,1039,1058]" italics="true" pageId="7" pageNumber="7">82</emphasis>
,
<emphasis id="289CEAF2FA7D8A1259E0FBB5FE17A15E" bold="true" box="[424,446,1039,1058]" italics="true" pageId="7" pageNumber="7">84</emphasis>
) (
<figureCitation id="82D32A65FA7D8A12599EFBB5FDA5A15E" box="[470,524,1039,1058]" captionStart="Fig. 6" captionStartId="8.[573,604,771,788]" captionTargetBox="[837,1482,762,1761]" captionTargetId="figure@8.[827,1486,758,1765]" captionTargetPageId="8" captionText="Fig. 6. Elbow and wrist morphology. The most diagnostic features of the elbow and wrist joints of Pliobates cataloniae gen. et sp. nov. (IPS 58443), denoted by arrows in drawings of the distal humerus, proximal radius, and distal ulna, are shown with those of selected extant and extinct anthropoids for comparison. (A to D) Anterior (top) and distal (bottom) views of the distal humerus in P. cata- loniae (A), Epipliopithecus vindobonensis Individual I [(B), reversed], Dendro- pithecus? sp. KNM MO 17022 A (C) (MO, Moruorot), and Hylobates moloch (D). (E to H) Views perpendicular to the radial tuberosity (top) and proximal view (bottom) of the proximal radius in P. cataloniae (E), E. vindobonensis Individual I (F), Simiolus enjiessi KNM MO 63 [(G), reversed)], and H. moloch (H). (I to M) Medial (top) and distal (bottom) views of the distal ulna in P. cataloniae (I), E. vindobonensis Indi- vidual I (J), H. moloch (K), Ateles paniscus (L), and Cercopithecus aethiops (M). 1, absence of entepicondylar foramen; 2, absence of capitular tail; 3, lack of spool-shaped trochlea; 4, well-developed beveled surface for the zona conoidea; 5, small and flat area in the radial head; 6, ulnar fovea; 7, two-faceted, expanded semilunar articular surface in the ulnar head. Specimens are shown as if from the left side and are not to scale. [Artwork by M. Palmero" httpUri="https://zenodo.org/record/269430/files/figure.png" pageId="7" pageNumber="7">Fig. 6</figureCitation>
), including: in the humerus, the lack of capitular tail [present in
<taxonomicName id="DDE84D63FA7D8A1258A2FBFEFEDEA12B" authority="Zapfe &amp; Hürzeler, 1957" authorityName="Zapfe &amp; Hürzeler" authorityYear="1957" box="[234,375,1092,1111]" class="Mammalia" genus="Epipliopithecus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="7" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="289CEAF2FA7D8A1258A2FBFEFEDEA12B" bold="true" box="[234,375,1092,1111]" italics="true" pageId="7" pageNumber="7">Epipliopithecus</emphasis>
</taxonomicName>
, dendropithecids, and cercopithecoids (
<emphasis id="289CEAF2FA7D8A125965FBE4FEEDA10D" bold="true" box="[301,324,1118,1137]" italics="true" pageId="7" pageNumber="7">84</emphasis>
)] and the moderately globulous (although not posterolaterally expanded) capitulum with a well-developed zona conoidea [lacking in
<taxonomicName id="DDE84D63FA7D8A125881FB14FEFFA1BD" box="[201,342,1198,1217]" pageId="7" pageNumber="7">
<emphasis id="289CEAF2FA7D8A125881FB14FEFFA1BD" bold="true" box="[201,342,1198,1217]" italics="true" pageId="7" pageNumber="7">Epipliopithecus</emphasis>
</taxonomicName>
and dendropithecids (
<emphasis id="289CEAF2FA7D8A125821FB73FF35A1A0" bold="true" box="[105,156,1225,1244]" italics="true" pageId="7" pageNumber="7">8184</emphasis>
)]; and, in the radius, the only slightly tilted and almost circular radial head with a small and flat area, a reduced lateral lip, and a beveled surface for the humeral zona conoidea.
<taxonomicName id="DDE84D63FA7D8A12598FFAA2FDB1A057" box="[455,536,1304,1323]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="7" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7D8A12598FFAA2FDB1A057" bold="true" box="[455,536,1304,1323]" italics="true" pageId="7" pageNumber="7">Pliobates</emphasis>
</taxonomicName>
also has a hominoid-like diarthrodial distal radioulnar joint (
<emphasis id="289CEAF2FA7D8A125884FAF7FEA9A01C" bold="true" box="[204,256,1357,1377]" italics="true" pageId="7" pageNumber="7">8587</emphasis>
), with a two-faceted expanded semilunar articulation in the ulnar head (
<figureCitation id="82D32A65FA7D8A125991FAD2FDA4A007" box="[473,525,1384,1403]" captionStart="Fig. 6" captionStartId="8.[573,604,771,788]" captionTargetBox="[837,1482,762,1761]" captionTargetId="figure@8.[827,1486,758,1765]" captionTargetPageId="8" captionText="Fig. 6. Elbow and wrist morphology. The most diagnostic features of the elbow and wrist joints of Pliobates cataloniae gen. et sp. nov. (IPS 58443), denoted by arrows in drawings of the distal humerus, proximal radius, and distal ulna, are shown with those of selected extant and extinct anthropoids for comparison. (A to D) Anterior (top) and distal (bottom) views of the distal humerus in P. cata- loniae (A), Epipliopithecus vindobonensis Individual I [(B), reversed], Dendro- pithecus? sp. KNM MO 17022 A (C) (MO, Moruorot), and Hylobates moloch (D). (E to H) Views perpendicular to the radial tuberosity (top) and proximal view (bottom) of the proximal radius in P. cataloniae (E), E. vindobonensis Individual I (F), Simiolus enjiessi KNM MO 63 [(G), reversed)], and H. moloch (H). (I to M) Medial (top) and distal (bottom) views of the distal ulna in P. cataloniae (I), E. vindobonensis Indi- vidual I (J), H. moloch (K), Ateles paniscus (L), and Cercopithecus aethiops (M). 1, absence of entepicondylar foramen; 2, absence of capitular tail; 3, lack of spool-shaped trochlea; 4, well-developed beveled surface for the zona conoidea; 5, small and flat area in the radial head; 6, ulnar fovea; 7, two-faceted, expanded semilunar articular surface in the ulnar head. Specimens are shown as if from the left side and are not to scale. [Artwork by M. Palmero" httpUri="https://zenodo.org/record/269430/files/figure.png" pageId="7" pageNumber="7">Fig.6</figureCitation>
). In this regard,
<taxonomicName id="DDE84D63FA7D8A1258BAFA39FEECA0EA" box="[242,325,1411,1430]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="7" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7D8A1258BAFA39FEECA0EA" bold="true" box="[242,325,1411,1430]" italics="true" pageId="7" pageNumber="7">Pliobates</emphasis>
</taxonomicName>
departs from
<taxonomicName id="DDE84D63FA7D8A125986FA39FF07A0CC" pageId="7" pageNumber="7">
<emphasis id="289CEAF2FA7D8A125986FA39FF07A0CC" bold="true" italics="true" pageId="7" pageNumber="7">Epipliopithecus</emphasis>
</taxonomicName>
, dendropithecids, and cercopithecoids (
<emphasis id="289CEAF2FA7D8A125821FA02FFD3A0B7" bold="true" box="[105,122,1464,1483]" italics="true" pageId="7" pageNumber="7">17</emphasis>
,
<emphasis id="289CEAF2FA7D8A1258CCFA02FF33A0B7" bold="true" box="[132,154,1464,1483]" italics="true" pageId="7" pageNumber="7">38</emphasis>
,
<emphasis id="289CEAF2FA7D8A1258EBFA02FF10A0B7" bold="true" box="[163,185,1464,1483]" italics="true" pageId="7" pageNumber="7">82</emphasis>
) and more closely resembles
<emphasis id="289CEAF2FA7D8A1259F7FA02FDB1A0B7" bold="true" box="[447,536,1464,1483]" italics="true" pageId="7" pageNumber="7">Proconsul</emphasis>
(
<emphasis id="289CEAF2FA7D8A125821FA68FFD6A099" bold="true" box="[105,127,1490,1509]" italics="true" pageId="7" pageNumber="7">88</emphasis>
), although the ulnar head is less extensive than in extant hominoids. In contrast with these derived features, the humeral shaft and humeroulnar joint are plesiomorphic: The former (
<figureCitation id="82D32A65FA7D8A1259AAF998FDB8A349" box="[482,529,1570,1589]" captionStart="Fig. 5" captionStartId="7.[573,604,1758,1775]" captionTargetBox="[590,1468,989,1726]" captionTargetId="figure@7.[1013,1046,134,1913]" captionTargetPageId="7" captionText="Fig. 5. Forelimb long bones. Shown are the humerus, radius, and ulna of the holotype (IPS 58443) of Pliobates cataloniae gen. et sp. nov. (A to E) Partial left humerus in medial (A), posterior (B), lateral (C), anterior (D), and distal (E) views. (F to K) Left radius in medial (F), posterior (G), lateral (H), anterior (I), proximal (J), and distal (K) views. (L to O) Proximal half of the left ulna in medial (L), posterior (M), lateral (N), and anterior (O) views. (P to T) Distal fragment of the left ulna in medial (P), posterior (Q), lateral (R), anterior (S), and distal (T) views." httpUri="https://zenodo.org/record/269429/files/figure.png" pageId="7" pageNumber="7">Fig. 5</figureCitation>
) is anteriorly straight and somewhat proximally retroflexed; the latter (
<figureCitation id="82D32A65FA7D8A125963F9EDFEF5A316" box="[299,348,1623,1642]" captionStart="Fig. 6" captionStartId="8.[573,604,771,788]" captionTargetBox="[837,1482,762,1761]" captionTargetId="figure@8.[827,1486,758,1765]" captionTargetPageId="8" captionText="Fig. 6. Elbow and wrist morphology. The most diagnostic features of the elbow and wrist joints of Pliobates cataloniae gen. et sp. nov. (IPS 58443), denoted by arrows in drawings of the distal humerus, proximal radius, and distal ulna, are shown with those of selected extant and extinct anthropoids for comparison. (A to D) Anterior (top) and distal (bottom) views of the distal humerus in P. cata- loniae (A), Epipliopithecus vindobonensis Individual I [(B), reversed], Dendro- pithecus? sp. KNM MO 17022 A (C) (MO, Moruorot), and Hylobates moloch (D). (E to H) Views perpendicular to the radial tuberosity (top) and proximal view (bottom) of the proximal radius in P. cataloniae (E), E. vindobonensis Individual I (F), Simiolus enjiessi KNM MO 63 [(G), reversed)], and H. moloch (H). (I to M) Medial (top) and distal (bottom) views of the distal ulna in P. cataloniae (I), E. vindobonensis Indi- vidual I (J), H. moloch (K), Ateles paniscus (L), and Cercopithecus aethiops (M). 1, absence of entepicondylar foramen; 2, absence of capitular tail; 3, lack of spool-shaped trochlea; 4, well-developed beveled surface for the zona conoidea; 5, small and flat area in the radial head; 6, ulnar fovea; 7, two-faceted, expanded semilunar articular surface in the ulnar head. Specimens are shown as if from the left side and are not to scale. [Artwork by M. Palmero" httpUri="https://zenodo.org/record/269430/files/figure.png" pageId="7" pageNumber="7">Fig. 6</figureCitation>
) lacks the stabilizing features of extant hominoids (
<emphasis id="289CEAF2FA7D8A125930F9C8FE27A3F9" bold="true" box="[376,398,1650,1669]" italics="true" pageId="7" pageNumber="7">83</emphasis>
,
<emphasis id="289CEAF2FA7D8A1259D2F9C8FE19A3F9" bold="true" box="[410,432,1650,1669]" italics="true" pageId="7" pageNumber="7">89</emphasis>
), as shown by the narrow ulnar trochlear notch without a median keel (in agreement with the absence of spooling and the poorly defined trochlear lateral keel in the humerus of
<taxonomicName id="DDE84D63FA7D8A12597AF966FE28A393" box="[306,385,1756,1775]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="7" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7D8A12597AF966FE28A393" bold="true" box="[306,385,1756,1775]" italics="true" pageId="7" pageNumber="7">Pliobates</emphasis>
</taxonomicName>
).
</paragraph>
<caption id="4E976668FA7D8A125A75F964FC22A20F" httpUri="https://zenodo.org/record/269429/files/figure.png" pageId="7" pageNumber="7" targetBox="[590,1468,989,1726]" targetPageId="7">
<paragraph id="1A5736E0FA7D8A125A75F964FC22A20F" blockId="7.[573,1486,1756,1908]" pageId="7" pageNumber="7">
<emphasis id="289CEAF2FA7D8A125A75F964FCE0A392" bold="true" box="[573,841,1758,1775]" pageId="7" pageNumber="7">Fig. 5. Forelimb long bones.</emphasis>
Shown are the humerus, radius, and ulna of the holotype (IPS58443) of
<taxonomicName id="DDE84D63FA7D8A125A1CF942FCACA275" box="[596,773,1784,1801]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="7" pageNumber="7" phylum="Chordata" rank="species" species="cataloniae">
<emphasis id="289CEAF2FA7D8A125A1CF942FCACA275" box="[596,773,1784,1801]" italics="true" pageId="7" pageNumber="7">Pliobates cataloniae</emphasis>
</taxonomicName>
gen. et sp. nov. (
<emphasis id="289CEAF2FA7D8A125BD7F942FC04A275" bold="true" box="[927,941,1784,1801]" pageId="7" pageNumber="7">A</emphasis>
to
<emphasis id="289CEAF2FA7D8A125B82F942FC7FA274" bold="true" box="[970,982,1784,1800]" pageId="7" pageNumber="7">E</emphasis>
) Partial left humerus in medial (A), posterior (B), lateral (C), anterior (D), and distal (E) views. (
<emphasis id="289CEAF2FA7D8A125BC7F8A9FC32A25F" bold="true" box="[911,923,1811,1827]" pageId="7" pageNumber="7">F</emphasis>
to
<emphasis id="289CEAF2FA7D8A125BF0F8A9FC6CA25F" bold="true" box="[952,965,1811,1827]" pageId="7" pageNumber="7">K</emphasis>
) Left radius in medial (F), posterior (G), lateral (H), anterior (I), proximal (J), and distal (K) views. (
<emphasis id="289CEAF2FA7D8A125B34F894FC21A242" bold="true" box="[892,904,1838,1854]" pageId="7" pageNumber="7">L</emphasis>
to
<emphasis id="289CEAF2FA7D8A125BE9F897FC19A242" bold="true" box="[929,944,1837,1854]" pageId="7" pageNumber="7">O</emphasis>
) Proximal half of the left ulna in medial (L), posterior (M), lateral (N), and anterior (O) views. (
<emphasis id="289CEAF2FA7D8A125B77F8F2FCE5A224" bold="true" box="[831,844,1864,1880]" pageId="7" pageNumber="7">P</emphasis>
to
<emphasis id="289CEAF2FA7D8A125B23F8F2FCDEA224" bold="true" box="[875,887,1864,1880]" pageId="7" pageNumber="7">T</emphasis>
) Distal fragment of the left ulna in medial (P), posterior (Q), lateral (R), anterior (S), and distal (T) views.
</paragraph>
</caption>
<paragraph id="1A5736E0FA7D8A125830F94DFABAA4D8" blockId="7.[98,537,817,1908]" lastBlockId="7.[1048,1487,135,925]" pageId="7" pageNumber="7">
Humeral torsion in
<taxonomicName id="DDE84D63FA7D8A12597DF94DFE2FA276" box="[309,390,1783,1802]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="7" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7D8A12597DF94DFE2FA276" bold="true" box="[309,390,1783,1802]" italics="true" pageId="7" pageNumber="7">Pliobates</emphasis>
</taxonomicName>
is estimated at 101°, irrespective of the method employed (based on the posterior buttress for the humeral head or the bisector of the bicipital groove), with a confidence interval spanning 95.7° to 106.3° [based on the prediction error (5.23%) for the bicipital groove method (
<emphasis id="289CEAF2FA7D8A125A92FF18FD58A5C9" bold="true" box="[730,753,162,181]" italics="true" pageId="7" pageNumber="7">50</emphasis>
)]. This degree of torsion is moderate, higher than that estimated for
<emphasis id="289CEAF2FA7D8A125B84FF07FD6EA596" bold="true" italics="true" pageId="7" pageNumber="7">Proconsul heseloni</emphasis>
(92°), but comparable to estimates for
<emphasis id="289CEAF2FA7D8A125A11FF48FCA2A479" bold="true" box="[601,779,242,261]" italics="true" pageId="7" pageNumber="7">Dryopithecus fontani</emphasis>
(102°) and
<emphasis id="289CEAF2FA7D8A125B23FF48FD33A463" bold="true" italics="true" pageId="7" pageNumber="7">Dendropithecus macinnesi</emphasis>
(103.5°), and only slightly below the value estimated for
<taxonomicName id="DDE84D63FA7D8A125AA0FE9DFC5BA446" box="[744,1010,295,314]" pageId="7" pageNumber="7">
<emphasis id="289CEAF2FA7D8A125AA0FE9DFC5BA446" bold="true" box="[744,1010,295,314]" italics="true" pageId="7" pageNumber="7">Epipliopithecus vindobonensis</emphasis>
</taxonomicName>
(109°). The humeral torsion of
<taxonomicName id="DDE84D63FA7D8A125B15FEFBFC07A428" box="[861,942,321,340]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="7" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7D8A125B15FEFBFC07A428" bold="true" box="[861,942,321,340]" italics="true" pageId="7" pageNumber="7">Pliobates</emphasis>
</taxonomicName>
is thus most comparable to that of non-atelid platyrrhines and lower than that of
<taxonomicName id="DDE84D63FA7D8A125B5EFECCFCE3A4F5" box="[790,842,374,393]" class="Mammalia" family="Atelidae" genus="Ateles" kingdom="Animalia" order="Primates" pageId="7" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="289CEAF2FA7D8A125B5EFECCFCE3A4F5" bold="true" box="[790,842,374,393]" italics="true" pageId="7" pageNumber="7">Ateles</emphasis>
</taxonomicName>
and extant hominoids, especially African great apes and humans [although the high degree of humeral torsion of extant hominoids is related to increased mobility at the glenohumeral joint, the higher values of great apes and humans appear related to knucklewalking and enhanced manipulation, respectively, rather than suspensory behaviors (
<emphasis id="289CEAF2FA7D8A125BC8FD8AFC3FA73F" bold="true" box="[896,918,560,579]" italics="true" pageId="7" pageNumber="7">50</emphasis>
)]. In contrast to the moderate humeral torsion, the forelimb of
<taxonomicName id="DDE84D63FA7D8A125AC2FDDCFD72A705" box="[650,731,614,633]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="7" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7D8A125AC2FDDCFD72A705" bold="true" box="[650,731,614,633]" italics="true" pageId="7" pageNumber="7">Pliobates</emphasis>
</taxonomicName>
appears somewhat elongated relative to its body size (fig. S5). Allometric computations of relative forelimb length in fossils (residuals are given in table S14) must be considered with caution, because they are dependent on the accuracy of body-size estimates. However, the forelimb of
<taxonomicName id="DDE84D63FA7D8A125A83FCBFFCB5A664" box="[715,796,773,792]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="7" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7D8A125A83FCBFFCB5A664" bold="true" box="[715,796,773,792]" italics="true" pageId="7" pageNumber="7">Pliobates</emphasis>
</taxonomicName>
(based on our BM estimate of 4.5 kg) appears more elongated than that of
<taxonomicName id="DDE84D63FA7D8A125A11FC80FD45A631" box="[601,748,826,845]" pageId="7" pageNumber="7">
<emphasis id="289CEAF2FA7D8A125A11FC80FD45A631" bold="true" box="[601,748,826,845]" italics="true" pageId="7" pageNumber="7">Epipliopithecus</emphasis>
</taxonomicName>
(based on our estimate of 11.5 kg). The latter taxon, contrary to previous assertions (
<emphasis id="289CEAF2FA7D8A125AEFFCD5FD13A6FE" bold="true" box="[679,698,879,898]" italics="true" pageId="7" pageNumber="7">81</emphasis>
,
<emphasis id="289CEAF2FA7D8A125A8FFCD5FD74A6FE" bold="true" box="[711,733,879,898]" italics="true" pageId="7" pageNumber="7">83</emphasis>
), has the generalized proportions of quadrupedal monkeys.
<taxonomicName id="DDE84D63FA7D8A125B11FC30FC03A6E1" box="[857,938,906,925]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="7" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7D8A125B11FC30FC03A6E1" bold="true" box="[857,938,906,925]" italics="true" pageId="7" pageNumber="7">Pliobates</emphasis>
</taxonomicName>
, in contrast, has a forelimb elongation similar to that of female orangutans and
<taxonomicName id="DDE84D63FA7D8A125CBDFF18FAF5A5C9" box="[1269,1372,162,181]" class="Mammalia" family="Atelidae" genus="Brachyteles" kingdom="Animalia" order="Primates" pageId="7" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="289CEAF2FA7D8A125CBDFF18FAF5A5C9" bold="true" box="[1269,1372,162,181]" italics="true" pageId="7" pageNumber="7">Brachyteles</emphasis>
</taxonomicName>
, although it is less extreme than in
<taxonomicName id="DDE84D63FA7D8A125CAFFF07FAB2A5AC" box="[1255,1307,189,208]" class="Mammalia" family="Atelidae" genus="Ateles" kingdom="Animalia" order="Primates" pageId="7" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="289CEAF2FA7D8A125CAFFF07FAB2A5AC" bold="true" box="[1255,1307,189,208]" italics="true" pageId="7" pageNumber="7">Ateles</emphasis>
</taxonomicName>
and especially than in hylobatids. The same pattern holds when the humerus and radius are analyzed separately, although in
<taxonomicName id="DDE84D63FA7D8A125CC7FEB6FB49A463" box="[1167,1248,268,287]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="7" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7D8A125CC7FEB6FB49A463" bold="true" box="[1167,1248,268,287]" italics="true" pageId="7" pageNumber="7">Pliobates</emphasis>
</taxonomicName>
, relative length is somewhat higher for the radius than for the humerus.
<taxonomicName id="DDE84D63FA7D8A125C50FEFBFBC0A428" box="[1048,1129,321,340]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="7" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7D8A125C50FEFBFBC0A428" bold="true" box="[1048,1129,321,340]" italics="true" pageId="7" pageNumber="7">Pliobates</emphasis>
</taxonomicName>
further displays a high arm angle (8°), which is considerably greater than the average in most anthropoids, except
<taxonomicName id="DDE84D63FA7D8A125CB6FECCFAFEA4F5" box="[1278,1367,374,393]" class="Mammalia" family="Hylobatidae" genus="Hylobates" kingdom="Animalia" order="Primates" pageId="7" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="289CEAF2FA7D8A125CB6FECCFAFEA4F5" bold="true" box="[1278,1367,374,393]" italics="true" pageId="7" pageNumber="7">Hylobates</emphasis>
</taxonomicName>
(9.8°),
<taxonomicName id="DDE84D63FA7D8A125DDEFECCFA64A4F5" box="[1430,1485,374,393]" class="Mammalia" family="Hominidae" genus="Pongo" kingdom="Animalia" order="Primates" pageId="7" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="289CEAF2FA7D8A125DDEFECCFA64A4F5" bold="true" box="[1430,1485,374,393]" italics="true" pageId="7" pageNumber="7">Pongo</emphasis>
</taxonomicName>
(6.3°), and
<taxonomicName id="DDE84D63FA7D8A125C35FE2BFB18A4D8" box="[1149,1201,401,420]" class="Mammalia" family="Atelidae" genus="Ateles" kingdom="Animalia" order="Primates" pageId="7" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="289CEAF2FA7D8A125C35FE2BFB18A4D8" bold="true" box="[1149,1201,401,420]" italics="true" pageId="7" pageNumber="7">Ateles</emphasis>
</taxonomicName>
(6.5°) (
<emphasis id="289CEAF2FA7D8A125CBCFE2BFAAFA4D8" bold="true" box="[1268,1286,401,420]" italics="true" pageId="7" pageNumber="7">51</emphasis>
).
</paragraph>
<paragraph id="1A5736E0FA7D8A1D5C65FE16FE7BA17B" blockId="7.[1048,1487,135,925]" lastBlockId="8.[98,536,135,1908]" lastPageId="8" lastPageNumber="8" pageId="7" pageNumber="7">
The ulnocarpal articulation of
<taxonomicName id="DDE84D63FA7D8A125D17FE16FA1DA4C3" box="[1375,1460,428,447]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="7" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7D8A125D17FE16FA1DA4C3" bold="true" box="[1375,1460,428,447]" italics="true" pageId="7" pageNumber="7">Pliobates</emphasis>
</taxonomicName>
is completely different from that of
<taxonomicName id="DDE84D63FA7D8A125D28FE7CFB9AA488" pageId="7" pageNumber="7">
<emphasis id="289CEAF2FA7D8A125D28FE7CFB9AA488" bold="true" italics="true" pageId="7" pageNumber="7">Epipliopithecus</emphasis>
</taxonomicName>
and dendropithecids (
<emphasis id="289CEAF2FA7D8A125D4FFE5BFAB0A488" bold="true" box="[1287,1305,481,500]" italics="true" pageId="7" pageNumber="7">17</emphasis>
,
<emphasis id="289CEAF2FA7D8A125D6AFE5BFA90A488" bold="true" box="[1314,1337,481,500]" italics="true" pageId="7" pageNumber="7">38</emphasis>
,
<emphasis id="289CEAF2FA7D8A125D0BFE5BFAF3A488" bold="true" box="[1347,1370,481,500]" italics="true" pageId="7" pageNumber="7">90</emphasis>
), including a partially developed ulnar fovea (
<figureCitation id="82D32A65FA7D8A125D7CFE41FACCA772" box="[1332,1381,507,526]" captionStart="Fig. 6" captionStartId="8.[573,604,771,788]" captionTargetBox="[837,1482,762,1761]" captionTargetId="figure@8.[827,1486,758,1765]" captionTargetPageId="8" captionText="Fig. 6. Elbow and wrist morphology. The most diagnostic features of the elbow and wrist joints of Pliobates cataloniae gen. et sp. nov. (IPS 58443), denoted by arrows in drawings of the distal humerus, proximal radius, and distal ulna, are shown with those of selected extant and extinct anthropoids for comparison. (A to D) Anterior (top) and distal (bottom) views of the distal humerus in P. cata- loniae (A), Epipliopithecus vindobonensis Individual I [(B), reversed], Dendro- pithecus? sp. KNM MO 17022 A (C) (MO, Moruorot), and Hylobates moloch (D). (E to H) Views perpendicular to the radial tuberosity (top) and proximal view (bottom) of the proximal radius in P. cataloniae (E), E. vindobonensis Individual I (F), Simiolus enjiessi KNM MO 63 [(G), reversed)], and H. moloch (H). (I to M) Medial (top) and distal (bottom) views of the distal ulna in P. cataloniae (I), E. vindobonensis Indi- vidual I (J), H. moloch (K), Ateles paniscus (L), and Cercopithecus aethiops (M). 1, absence of entepicondylar foramen; 2, absence of capitular tail; 3, lack of spool-shaped trochlea; 4, well-developed beveled surface for the zona conoidea; 5, small and flat area in the radial head; 6, ulnar fovea; 7, two-faceted, expanded semilunar articular surface in the ulnar head. Specimens are shown as if from the left side and are not to scale. [Artwork by M. Palmero" httpUri="https://zenodo.org/record/269430/files/figure.png" pageId="7" pageNumber="7">Fig. 6</figureCitation>
), which, in extant hominoids, is the attachment area of the triangular disc ligament and the intra-articular meniscus (
<emphasis id="289CEAF2FA7D8A125C36FDF1FB1DA722" bold="true" box="[1150,1204,587,606]" italics="true" pageId="7" pageNumber="7">8587</emphasis>
). The ulnar styloid process is relatively long and slender, with no discernible articular surfaces for the pisiform or triquetrum. This agrees with the lack of an articular facet for the styloid process on the pisiform, like extant hominoids but unlike monkeys and
<emphasis id="289CEAF2FA7D8A125D39FD6AFA67A79F" bold="true" box="[1393,1486,720,739]" italics="true" pageId="7" pageNumber="7">Proconsul</emphasis>
(
<emphasis id="289CEAF2FA7D8A125C57FD50FB9BA781" bold="true" box="[1055,1074,746,765]" italics="true" pageId="7" pageNumber="7">91</emphasis>
). However, in contrast to
<emphasis id="289CEAF2FA7D8A125D5EFD50FA08A781" bold="true" box="[1302,1441,746,765]" italics="true" pageId="7" pageNumber="7">Pierolapithecus</emphasis>
(
<emphasis id="289CEAF2FA7D8A125DE4FD50FA6BA781" bold="true" box="[1452,1474,746,765]" italics="true" pageId="7" pageNumber="7">92</emphasis>
), the triquetrum of
<taxonomicName id="DDE84D63FA7D8A125C80FCBFFAB4A664" box="[1224,1309,773,792]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="7" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA7D8A125C80FCBFFAB4A664" bold="true" box="[1224,1309,773,792]" italics="true" pageId="7" pageNumber="7">Pliobates</emphasis>
</taxonomicName>
shows a proximal articular facet, which is more developed than that present in hylobatids and sometimes
<taxonomicName id="DDE84D63FA7D8A125DE1FC80FA64A631" box="[1449,1485,826,845]" class="Mammalia" family="Hominidae" genus="Pan" kingdom="Animalia" order="Primates" pageId="7" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="289CEAF2FA7D8A125DE1FC80FA64A631" bold="true" box="[1449,1485,826,845]" italics="true" pageId="7" pageNumber="7">Pan</emphasis>
</taxonomicName>
(
<emphasis id="289CEAF2FA7D8A125C57FCEFFB9BA614" bold="true" box="[1055,1074,853,872]" italics="true" pageId="7" pageNumber="7">51</emphasis>
,
<emphasis id="289CEAF2FA7D8A125C09FCEFFBFFA614" bold="true" box="[1089,1110,853,872]" italics="true" pageId="7" pageNumber="7">87</emphasis>
) but less developed than in monkeys. This suggests that ulnotriquetral contact might have been reduced by some kind of intra-articular tissue, similarly to some
<taxonomicName id="DDE84D63FA728A1D5973FF3DFEC7A5E6" box="[315,366,135,154]" class="Mammalia" family="Atelidae" genus="Ateles" kingdom="Animalia" order="Primates" pageId="8" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis id="289CEAF2FA728A1D5973FF3DFEC7A5E6" bold="true" box="[315,366,135,154]" italics="true" pageId="8" pageNumber="8">Ateles</emphasis>
</taxonomicName>
species (
<emphasis id="289CEAF2FA728A1D59F4FF3DFE78A5E6" bold="true" box="[444,465,135,154]" italics="true" pageId="8" pageNumber="8">93</emphasis>
). Moreover, as in apes, the triquetrum of
<taxonomicName id="DDE84D63FA728A1D59E6FF18FDAAA5C9" box="[430,515,162,181]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="8" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA728A1D59E6FF18FDAAA5C9" bold="true" box="[430,515,162,181]" italics="true" pageId="8" pageNumber="8">Pliobates</emphasis>
</taxonomicName>
is small relative to hamate size (fig. S6), indicating a reduced loading on the ulnar side of the wrist. However, as in monkeys and
<emphasis id="289CEAF2FA728A1D59FEFF48FDBAA479" bold="true" box="[438,531,242,261]" italics="true" pageId="8" pageNumber="8">Proconsul</emphasis>
, the triquetrum of
<taxonomicName id="DDE84D63FA728A1D5943FEB6FEC9A463" box="[267,352,268,287]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="8" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA728A1D5943FEB6FEC9A463" bold="true" box="[267,352,268,287]" italics="true" pageId="8" pageNumber="8">Pliobates</emphasis>
</taxonomicName>
differs from that of extant apes and
<emphasis id="289CEAF2FA728A1D58B1FE9DFE2DA446" bold="true" box="[249,388,295,314]" italics="true" pageId="8" pageNumber="8">Pierolapithecus</emphasis>
(
<emphasis id="289CEAF2FA728A1D59D8FE9DFE0FA446" bold="true" box="[400,422,295,314]" italics="true" pageId="8" pageNumber="8">92</emphasis>
) by possessing a proximally protruding beak-like process (
<figureCitation id="82D32A65FA728A1D5820FEE6FF3FA413" box="[104,150,348,367]" captionStart="Fig. 7" captionStartId="9.[98,129,138,155]" captionTargetBox="[354,1010,134,901]" captionTargetId="figure@9.[352,1011,134,902]" captionTargetPageId="9" captionText="Fig. 7. Carpal bones. Line drawings of carpal bones in Pliobates cata- loniae gen. et sp. nov. (IPS 58443) are shown with those of selected anthropoid genera for comparison. (A to E) Left capitate, in radial (top) and proximal (bottom) views, of Cer- copithecus aethiops (A), Ateles paniscus (B), Pierolapithecus cat- alaunicus (C), Hylobates lar (D), and P. cataloniae (E); gray shading denotes articular areas for the second metacar- pals, and cross-hatching denotes those for the third metacarpal. (F to J) Left hamate, in radial (top) and ulnar (bottom) views, of C. aethiops (F), A. paniscus (G), Pi. catalaunicus (H), H. lar (I), and P. catalo- niae (J). (K to O) Left triquetrum, in proximo- medial (top) and distal (bottom) views, of C. aethiops (K), A. paniscus (L), Pi. catalaunicus (M), H. lar (N), and P. cataloniae (O). Drawings are not to scale." httpUri="https://zenodo.org/record/269431/files/figure.png" pageId="8" pageNumber="8">Fig. 7</figureCitation>
). The hamate of
<taxonomicName id="DDE84D63FA728A1D5962FEE6FED2A413" box="[298,379,348,367]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="8" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA728A1D5962FEE6FED2A413" bold="true" box="[298,379,348,367]" italics="true" pageId="8" pageNumber="8">Pliobates</emphasis>
</taxonomicName>
is “Miocene ape like” (
<emphasis id="289CEAF2FA728A1D58DFFECCFF03A4F5" bold="true" box="[151,170,374,393]" italics="true" pageId="8" pageNumber="8">91</emphasis>
,
<emphasis id="289CEAF2FA728A1D58FDFECCFF62A4F5" bold="true" box="[181,203,374,393]" italics="true" pageId="8" pageNumber="8">92</emphasis>
), although it more closely resembles that of hylobatids by possessing a dorsopalmarly narrow and proximodistally long triquetral articular surface that is proximally globular, as well as a distally projecting hamulus (
<figureCitation id="82D32A65FA728A1D59CFFE5BFE10A488" box="[391,441,481,500]" captionStart="Fig. 7" captionStartId="9.[98,129,138,155]" captionTargetBox="[354,1010,134,901]" captionTargetId="figure@9.[352,1011,134,902]" captionTargetPageId="9" captionText="Fig. 7. Carpal bones. Line drawings of carpal bones in Pliobates cata- loniae gen. et sp. nov. (IPS 58443) are shown with those of selected anthropoid genera for comparison. (A to E) Left capitate, in radial (top) and proximal (bottom) views, of Cer- copithecus aethiops (A), Ateles paniscus (B), Pierolapithecus cat- alaunicus (C), Hylobates lar (D), and P. cataloniae (E); gray shading denotes articular areas for the second metacar- pals, and cross-hatching denotes those for the third metacarpal. (F to J) Left hamate, in radial (top) and ulnar (bottom) views, of C. aethiops (F), A. paniscus (G), Pi. catalaunicus (H), H. lar (I), and P. catalo- niae (J). (K to O) Left triquetrum, in proximo- medial (top) and distal (bottom) views, of C. aethiops (K), A. paniscus (L), Pi. catalaunicus (M), H. lar (N), and P. cataloniae (O). Drawings are not to scale." httpUri="https://zenodo.org/record/269431/files/figure.png" pageId="8" pageNumber="8">Fig. 7</figureCitation>
).
<taxonomicName id="DDE84D63FA728A1D598FFE5BFDB1A488" box="[455,536,481,500]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="8" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA728A1D598FFE5BFDB1A488" bold="true" box="[455,536,481,500]" italics="true" pageId="8" pageNumber="8">Pliobates</emphasis>
</taxonomicName>
further resembles hylobatids and
<taxonomicName id="DDE84D63FA728A1D59C2FE41FE14A772" box="[394,445,507,526]" class="Mammalia" family="Atelidae" genus="Ateles" kingdom="Animalia" order="Primates" pageId="8" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis id="289CEAF2FA728A1D59C2FE41FE14A772" bold="true" box="[394,445,507,526]" italics="true" pageId="8" pageNumber="8">Ateles</emphasis>
</taxonomicName>
by having an oblong and mediolaterally narrow capitate head that, like the facet for the hamate, is proximodistally aligned. This morphology contrasts with the more globulous, wider, and ulnarly inclined capitate head of other catarrhines, including
<emphasis id="289CEAF2FA728A1D582AFD21FF12A7D2" bold="true" box="[98,187,667,686]" italics="true" pageId="8" pageNumber="8">Proconsul</emphasis>
(
<emphasis id="289CEAF2FA728A1D5883FD21FF48A7D2" bold="true" box="[203,225,667,686]" italics="true" pageId="8" pageNumber="8">88</emphasis>
,
<emphasis id="289CEAF2FA728A1D58A7FD21FEABA7D2" bold="true" box="[239,258,667,686]" italics="true" pageId="8" pageNumber="8">91</emphasis>
) and
<emphasis id="289CEAF2FA728A1D5976FD21FE60A7D2" bold="true" box="[318,457,667,686]" italics="true" pageId="8" pageNumber="8">Pierolapithecus</emphasis>
(
<emphasis id="289CEAF2FA728A1D5991FD21FE46A7D2" bold="true" box="[473,495,667,686]" italics="true" pageId="8" pageNumber="8">92</emphasis>
); in
<taxonomicName id="DDE84D63FA728A1D582AFD0FFF1EA7B4" box="[98,183,693,712]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="8" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA728A1D582AFD0FFF1EA7B4" bold="true" box="[98,183,693,712]" italics="true" pageId="8" pageNumber="8">Pliobates</emphasis>
</taxonomicName>
, though, it is not radially inclined, as it is in hylobatids. Moreover, the capitate facet for the second metacarpal is divided by a deep ligamentary notch (
<figureCitation id="82D32A65FA728A1D58B1FCBFFE87A664" box="[249,302,773,792]" captionStart="Fig. 7" captionStartId="9.[98,129,138,155]" captionTargetBox="[354,1010,134,901]" captionTargetId="figure@9.[352,1011,134,902]" captionTargetPageId="9" captionText="Fig. 7. Carpal bones. Line drawings of carpal bones in Pliobates cata- loniae gen. et sp. nov. (IPS 58443) are shown with those of selected anthropoid genera for comparison. (A to E) Left capitate, in radial (top) and proximal (bottom) views, of Cer- copithecus aethiops (A), Ateles paniscus (B), Pierolapithecus cat- alaunicus (C), Hylobates lar (D), and P. cataloniae (E); gray shading denotes articular areas for the second metacar- pals, and cross-hatching denotes those for the third metacarpal. (F to J) Left hamate, in radial (top) and ulnar (bottom) views, of C. aethiops (F), A. paniscus (G), Pi. catalaunicus (H), H. lar (I), and P. catalo- niae (J). (K to O) Left triquetrum, in proximo- medial (top) and distal (bottom) views, of C. aethiops (K), A. paniscus (L), Pi. catalaunicus (M), H. lar (N), and P. cataloniae (O). Drawings are not to scale." httpUri="https://zenodo.org/record/269431/files/figure.png" pageId="8" pageNumber="8">Fig. 7</figureCitation>
), as in extant hominoids and
<emphasis id="289CEAF2FA728A1D58C4FC9AFEBEA64F" bold="true" box="[140,279,800,819]" italics="true" pageId="8" pageNumber="8">Pierolapithecus</emphasis>
(
<emphasis id="289CEAF2FA728A1D596DFC9AFE92A64F" bold="true" box="[293,315,800,819]" italics="true" pageId="8" pageNumber="8">92</emphasis>
) but not in other catarrhines (including
<emphasis id="289CEAF2FA728A1D594EFC80FEF6A631" bold="true" box="[262,351,826,845]" italics="true" pageId="8" pageNumber="8">Proconsul</emphasis>
), in which the facet for the second metacarpal is dorsopalmarly continuous and occupies the whole lateral aspect of the capitate (
<figureCitation id="82D32A65FA728A1D58BAFC30FE8DA6E1" box="[242,292,906,925]" captionStart="Fig. 7" captionStartId="9.[98,129,138,155]" captionTargetBox="[354,1010,134,901]" captionTargetId="figure@9.[352,1011,134,902]" captionTargetPageId="9" captionText="Fig. 7. Carpal bones. Line drawings of carpal bones in Pliobates cata- loniae gen. et sp. nov. (IPS 58443) are shown with those of selected anthropoid genera for comparison. (A to E) Left capitate, in radial (top) and proximal (bottom) views, of Cer- copithecus aethiops (A), Ateles paniscus (B), Pierolapithecus cat- alaunicus (C), Hylobates lar (D), and P. cataloniae (E); gray shading denotes articular areas for the second metacar- pals, and cross-hatching denotes those for the third metacarpal. (F to J) Left hamate, in radial (top) and ulnar (bottom) views, of C. aethiops (F), A. paniscus (G), Pi. catalaunicus (H), H. lar (I), and P. catalo- niae (J). (K to O) Left triquetrum, in proximo- medial (top) and distal (bottom) views, of C. aethiops (K), A. paniscus (L), Pi. catalaunicus (M), H. lar (N), and P. cataloniae (O). Drawings are not to scale." httpUri="https://zenodo.org/record/269431/files/figure.png" pageId="8" pageNumber="8">Fig. 7</figureCitation>
) (
<emphasis id="289CEAF2FA728A1D5970FC30FEE2A6E1" bold="true" box="[312,331,906,925]" italics="true" pageId="8" pageNumber="8">91</emphasis>
).
<taxonomicName id="DDE84D63FA728A1D5915FC30FE07A6E1" box="[349,430,906,925]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="8" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA728A1D5915FC30FE07A6E1" bold="true" box="[349,430,906,925]" italics="true" pageId="8" pageNumber="8">Pliobates</emphasis>
</taxonomicName>
has a complex articulation between the third metacarpal and capitate, as in extant apes but not in
<emphasis id="289CEAF2FA728A1D5989FC05FDBEA6AE" bold="true" box="[449,535,959,978]" italics="true" pageId="8" pageNumber="8">Proconsul</emphasis>
(
<emphasis id="289CEAF2FA728A1D5821FC60FFD5A691" bold="true" box="[105,124,986,1005]" italics="true" pageId="8" pageNumber="8">91</emphasis>
); however, as in
<emphasis id="289CEAF2FA728A1D595AFC60FE31A691" bold="true" box="[274,408,986,1005]" italics="true" pageId="8" pageNumber="8">Pierolapithecus</emphasis>
(
<emphasis id="289CEAF2FA728A1D59EDFC60FE13A691" bold="true" box="[421,442,986,1005]" italics="true" pageId="8" pageNumber="8">92</emphasis>
), the capitate of
<taxonomicName id="DDE84D63FA728A1D58E8FC4EFF47A17B" box="[160,238,1012,1031]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="8" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA728A1D58E8FC4EFF47A17B" bold="true" box="[160,238,1012,1031]" italics="true" pageId="8" pageNumber="8">Pliobates</emphasis>
</taxonomicName>
lacks a hook-like process.
</paragraph>
<caption id="4E976668FA728A1D5A75FCB9FA67A20C" httpUri="https://zenodo.org/record/269430/files/figure.png" pageId="8" pageNumber="8" targetBox="[837,1482,762,1761]" targetPageId="8">
<paragraph id="1A5736E0FA728A1D5A75FCB9FA67A20C" pageId="8" pageNumber="8">
Fig. 6. Elbow and wrist
<emphasis id="289CEAF2FA728A1D5A75FCA4FD1EA652" bold="true" box="[573,695,798,814]" pageId="8" pageNumber="8">morphology.</emphasis>
The most diagnostic features of the elbow and wrist joints of
<taxonomicName id="DDE84D63FA728A1D5A75FCD4FD43A603" box="[573,746,878,895]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="cataloniae">
<emphasis id="289CEAF2FA728A1D5A75FCD4FD43A603" box="[573,746,878,895]" italics="true" pageId="8" pageNumber="8">Pliobates cataloniae</emphasis>
</taxonomicName>
gen. et sp. nov. (IPS58443), denoted by arrows in drawings of the distal humerus, proximal radius, and distal ulna, are shown with those of selected extant and extinct anthropoids for comparison. (
<emphasis id="289CEAF2FA728A1D5AFEFBE7FD6DA112" bold="true" box="[694,708,1117,1134]" pageId="8" pageNumber="8">A</emphasis>
to
<emphasis id="289CEAF2FA728A1D5AA8FBE7FD47A111" bold="true" box="[736,750,1117,1133]" pageId="8" pageNumber="8">D</emphasis>
) Anterior (top) and distal (bottom) views of the distal humerus in
<emphasis id="289CEAF2FA728A1D5A91FB17FDD9A1A4" italics="true" pageId="8" pageNumber="8">P. cata- loniae</emphasis>
(A),
<taxonomicName id="DDE84D63FA728A1D5AD0FB7DFD11A18F" pageId="8" pageNumber="8">
<emphasis id="289CEAF2FA728A1D5AD0FB7DFD11A18F" italics="true" pageId="8" pageNumber="8">Epipliopithecus vindobonensis</emphasis>
</taxonomicName>
Individual I [(B), reversed],
<emphasis id="289CEAF2FA728A1D5A8FFB46FD2FA054" italics="true" pageId="8" pageNumber="8">Dendro- pithecus</emphasis>
? sp. KNM MO 17022A (C) (MO, Moruorot), and
<taxonomicName id="DDE84D63FA728A1D5A8CFAF6FDD4A00B" class="Mammalia" family="Hylobatidae" genus="Hylobates" kingdom="Animalia" order="Primates" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="moloch">
<emphasis id="289CEAF2FA728A1D5A8CFAF6FDD4A00B" italics="true" pageId="8" pageNumber="8">Hylobates moloch</emphasis>
</taxonomicName>
(D). (
<emphasis id="289CEAF2FA728A1D5AF9FADDFD14A00B" bold="true" box="[689,701,1383,1399]" pageId="8" pageNumber="8">E</emphasis>
to
<emphasis id="289CEAF2FA728A1D5A91FADDFD41A00B" bold="true" box="[729,744,1383,1399]" pageId="8" pageNumber="8">H</emphasis>
) Views perpendicular to the radial tuberosity (top) and proximal view (bottom) of the proximal radius in
<taxonomicName id="DDE84D63FA728A1D5AC5FA56FD52A080" box="[653,763,1515,1532]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="cataloniae">
<emphasis id="289CEAF2FA728A1D5AC5FA56FD52A080" box="[653,763,1515,1532]" italics="true" pageId="8" pageNumber="8">P. cataloniae</emphasis>
</taxonomicName>
(E),
<emphasis id="289CEAF2FA728A1D5A2AF9BCFD5BA36B" box="[610,754,1542,1559]" italics="true" pageId="8" pageNumber="8">E. vindobonensis</emphasis>
Individual I (F),
<emphasis id="289CEAF2FA728A1D5A8BF99AFDD5A330" italics="true" pageId="8" pageNumber="8">Simiolus enjiessi</emphasis>
KNM MO 63 [(G), reversed)], and
<taxonomicName id="DDE84D63FA728A1D5A75F9CBFD3CA3FD" box="[573,661,1648,1665]" class="Mammalia" family="Hylobatidae" genus="Hylobates" kingdom="Animalia" order="Primates" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="moloch">
<emphasis id="289CEAF2FA728A1D5A75F9CBFD3CA3FD" box="[573,661,1648,1665]" italics="true" pageId="8" pageNumber="8">H. moloch</emphasis>
</taxonomicName>
(H). (
<emphasis id="289CEAF2FA728A1D5A82F9CAFD79A3FC" bold="true" box="[714,720,1648,1664]" pageId="8" pageNumber="8">I</emphasis>
to
<emphasis id="289CEAF2FA728A1D5AA4F9CAFD57A3FC" bold="true" box="[748,766,1648,1664]" pageId="8" pageNumber="8">M</emphasis>
) Medial (top) and distal (bottom) views of the distal ulna in
<taxonomicName id="DDE84D63FA728A1D5AE5F97AFCB2A3AD" box="[685,795,1728,1745]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="cataloniae">
<emphasis id="289CEAF2FA728A1D5AE5F97AFCB2A3AD" box="[685,795,1728,1745]" italics="true" pageId="8" pageNumber="8">P. cataloniae</emphasis>
</taxonomicName>
(I),
<emphasis id="289CEAF2FA728A1D5A13F961FD45A397" box="[603,748,1754,1771]" italics="true" pageId="8" pageNumber="8">E. vindobonensis</emphasis>
Indi- vidual I (J),
<taxonomicName id="DDE84D63FA728A1D5AD7F94CFD5FA27A" box="[671,758,1781,1798]" class="Mammalia" family="Hylobatidae" genus="Hylobates" kingdom="Animalia" order="Primates" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="moloch">
<emphasis id="289CEAF2FA728A1D5AD7F94CFD5FA27A" box="[671,758,1781,1798]" italics="true" pageId="8" pageNumber="8">H. moloch</emphasis>
</taxonomicName>
(K),
<taxonomicName id="DDE84D63FA728A1D5A75F8AAFD6AA25C" box="[573,707,1808,1825]" class="Mammalia" family="Atelidae" genus="Ateles" kingdom="Animalia" order="Primates" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="paniscus">
<emphasis id="289CEAF2FA728A1D5A75F8AAFD6AA25C" box="[573,707,1808,1825]" italics="true" pageId="8" pageNumber="8">Ateles paniscus</emphasis>
</taxonomicName>
(L), and
<taxonomicName id="DDE84D63FA728A1D5B5AF8AAFC73A25D" box="[786,986,1808,1825]" class="Mammalia" family="Cercopithecidae" genus="Cercopithecus" kingdom="Animalia" order="Primates" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="aethiops">
<emphasis id="289CEAF2FA728A1D5B5AF8AAFC73A25D" box="[786,986,1808,1825]" italics="true" pageId="8" pageNumber="8">Cercopithecus aethiops</emphasis>
</taxonomicName>
(M). 1, absence of entepicondylar foramen; 2, absence of capitular tail; 3, lack of spool-shaped trochlea; 4, well-developed beveled surface for the zona conoidea; 5, small and flat area in the radial head; 6, ulnar fovea; 7, two-faceted, expanded semilunar articular surface in the ulnar head. Specimens are shown as if from the left side and are not to scale. [Artwork by M. Palmero]
</paragraph>
</caption>
<paragraph id="1A5736E0FA728A1D5830FBB5FC1AA479" blockId="8.[98,536,135,1908]" lastBlockId="8.[573,1011,135,261]" pageId="8" pageNumber="8">
Regarding positional behaviors, although the primitive morphology of the proximal humerus of
<taxonomicName id="DDE84D63FA728A1D5834FBFEFF64A12B" box="[124,205,1092,1111]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="8" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA728A1D5834FBFEFF64A12B" bold="true" box="[124,205,1092,1111]" italics="true" pageId="8" pageNumber="8">Pliobates</emphasis>
</taxonomicName>
is suggestive of generalized abovebranch quadrupedalism (
<emphasis id="289CEAF2FA728A1D591CFBE4FEC3A10D" bold="true" box="[340,362,1118,1137]" italics="true" pageId="8" pageNumber="8">94</emphasis>
), its overall postcranial body plan is more compatible with a locomotor repertoire that includes a large amount of cautious and eclectic climbing (
<emphasis id="289CEAF2FA728A1D59EEFB14FE13A1BD" bold="true" box="[422,442,1198,1217]" italics="true" pageId="8" pageNumber="8">87</emphasis>
,
<emphasis id="289CEAF2FA728A1D598FFB14FE75A1BD" bold="true" box="[455,476,1198,1217]" italics="true" pageId="8" pageNumber="8">95</emphasis>
). This inference is supported by the emphasis on pronation and supination capabilities, the reduced compressive forces transferred across the ulnar side of the wrist, and the important ulnar deviation and rotatory capabilities. It agrees with previous hypotheses on the original locomotor adaptations of hominoids (
<emphasis id="289CEAF2FA728A1D58ADFAD2FF50A007" bold="true" box="[229,249,1384,1403]" italics="true" pageId="8" pageNumber="8">95</emphasis>
,
<emphasis id="289CEAF2FA728A1D594CFAD2FEB0A007" bold="true" box="[260,281,1384,1403]" italics="true" pageId="8" pageNumber="8">96</emphasis>
) and with recent interpretations of
<emphasis id="289CEAF2FA728A1D58E3FA39FEA9A0EA" bold="true" box="[171,256,1411,1430]" italics="true" pageId="8" pageNumber="8">Proconsul</emphasis>
that similarly depict this taxon as an arboreal quadruped with adaptations for cautious climbing and clambering (
<emphasis id="289CEAF2FA728A1D59FFFA02FE63A0B7" bold="true" box="[439,458,1464,1483]" italics="true" pageId="8" pageNumber="8">12</emphasis>
,
<emphasis id="289CEAF2FA728A1D599EFA02FE45A0B7" bold="true" box="[470,492,1464,1483]" italics="true" pageId="8" pageNumber="8">88</emphasis>
), including an incipient distal radioulnar diarthrosis that (unlike in
<taxonomicName id="DDE84D63FA728A1D58B9FA57FEEBA37C" box="[241,322,1517,1536]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="8" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA728A1D58B9FA57FEEBA37C" bold="true" box="[241,322,1517,1536]" italics="true" pageId="8" pageNumber="8">Pliobates</emphasis>
</taxonomicName>
) is still associated to a nonretreated ulnar styloid process (
<emphasis id="289CEAF2FA728A1D59E5F9BDFE6DA366" bold="true" box="[429,452,1543,1562]" italics="true" pageId="8" pageNumber="8">88</emphasis>
). The reduced ulnocarpal articulation of
<taxonomicName id="DDE84D63FA728A1D59DDF998FE43A349" box="[405,490,1570,1589]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="8" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA728A1D59DDF998FE43A349" bold="true" box="[405,490,1570,1589]" italics="true" pageId="8" pageNumber="8">Pliobates</emphasis>
</taxonomicName>
thus more closely foreshadows the condition of extant hominoids, although to a lesser extent than in the stem great ape
<emphasis id="289CEAF2FA728A1D595BF9C8FE37A3F9" bold="true" box="[275,414,1650,1669]" italics="true" pageId="8" pageNumber="8">Pierolapithecus</emphasis>
(
<emphasis id="289CEAF2FA728A1D59E3F9C8FE68A3F9" bold="true" box="[427,449,1650,1669]" italics="true" pageId="8" pageNumber="8">92</emphasis>
), indicating a decreased emphasis on forearm use under weight-bearing conditions relative to
<emphasis id="289CEAF2FA728A1D59F3F91DFDB1A3C6" bold="true" box="[443,536,1703,1722]" italics="true" pageId="8" pageNumber="8">Proconsul</emphasis>
(
<emphasis id="289CEAF2FA728A1D5822F97BFF28A3A8" bold="true" box="[106,129,1729,1748]" italics="true" pageId="8" pageNumber="8">88</emphasis>
). Several characteristics of
<taxonomicName id="DDE84D63FA728A1D5936F97BFE66A3A8" box="[382,463,1729,1748]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="8" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA728A1D5936F97BFE66A3A8" bold="true" box="[382,463,1729,1748]" italics="true" pageId="8" pageNumber="8">Pliobates</emphasis>
</taxonomicName>
(particularly the elongated forearm, the high arm angle, and the laterally facing bicipital tuberosity) further suggest some degree of below-branch forelimbdominated suspensory behaviors (
<emphasis id="289CEAF2FA728A1D59D6F896FE19A243" bold="true" box="[414,432,1836,1855]" italics="true" pageId="8" pageNumber="8">51</emphasis>
). However, the lack of hominoid-like elbow-stabilizing features in the humeroulnar joint (
<emphasis id="289CEAF2FA728A1D5903F8DBFEC9A208" bold="true" box="[331,352,1889,1908]" italics="true" pageId="8" pageNumber="8">83</emphasis>
,
<emphasis id="289CEAF2FA728A1D5923F8DBFE29A208" bold="true" box="[363,384,1889,1908]" italics="true" pageId="8" pageNumber="8">89</emphasis>
), the generalized metacarpophalangeal proportions, and the lack of marked phalangeal curvature suggest that
<taxonomicName id="DDE84D63FA728A1D5A75FF07FD23A5AC" box="[573,650,189,208]" class="Mammalia" family="Pliobatidae" genus="Pliobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Primates" pageId="8" pageNumber="4" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="289CEAF2FA728A1D5A75FF07FD23A5AC" bold="true" box="[573,650,189,208]" italics="true" pageId="8" pageNumber="8">Pliobates</emphasis>
</taxonomicName>
was not specifically adapted to perform the acrobatic suspensory behaviors (ricochetal brachiation) displayed by extant gibbons.
</paragraph>
</subSubSection>
</treatment>
</document>