375 lines
37 KiB
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375 lines
37 KiB
XML
<document ID-DOI="10.1002/evan.21929" ID-GBIF-Dataset="85a52735-5112-4407-b9bb-61dd3b03962d" ID-Zenodo-Dep="5627845" approvalRequired="1" approvalRequired_for_bibRefs="1" checkinTime="1635517063486" checkinUser="carolina" docAuthor="Roksandic, Mirjana, Radović, Predrag, Wu, Xiu-Jie & Bae, Christopher J." docDate="2022" docId="038C87AAFFEFFFA3FF5EFBF35514F916" docLanguage="en" docName="EvolAnthropol.31.1.20-29.pdf" docOrigin="Evolutionary Anthropology 31 (1)" docTitle="Homo bodoensis Roksandic & Radović & Wu & Bae 2021, sp. nov." docType="treatment" docVersion="9" lastPageNumber="6" masterDocId="FFB5FFD2FFEBFFA6FFFCFFC5517FFFE9" masterDocTitle="Resolving the “ muddle in the middle ”: The case for Homo bodoensis sp. nov." masterLastPageNumber="29" masterPageNumber="20" pageNumber="5" updateTime="1665099168891" updateUser="felipe">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>Resolving the “ muddle in the middle ”: The case for Homo bodoensis sp. nov.</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Roksandic, Mirjana</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Radović, Predrag</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Wu, Xiu-Jie</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Bae, Christopher J.</mods:namePart>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:relatedItem type="host">
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<mods:titleInfo>
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<mods:title>Evolutionary Anthropology</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2022</mods:date>
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<mods:detail type="pubDate">
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<mods:number>2021-10-28</mods:number>
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</mods:detail>
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<mods:detail type="volume">
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<mods:number>31</mods:number>
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</mods:detail>
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<mods:detail type="issue">
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<mods:number>1</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>20</mods:start>
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<mods:end>29</mods:end>
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</mods:extent>
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</mods:part>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">10.1002/evan.21929</mods:identifier>
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<mods:identifier type="GBIF-Dataset">85a52735-5112-4407-b9bb-61dd3b03962d</mods:identifier>
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<mods:identifier type="Zenodo-Dep">5627845</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.5634032" ID-GBIF-Taxon="189626279" ID-Zenodo-Dep="5634032" LSID="urn:lsid:zoobank.org:act:50AC3EA4-82E0-4AAD-BCDA-6DE6055888A7" httpUri="http://treatment.plazi.org/id/038C87AAFFEFFFA3FF5EFBF35514F916" lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5">
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<subSubSection box="[162,385,1078,1098]" pageId="4" pageNumber="5" type="nomenclature">
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<paragraph blockId="4.[162,506,867,1098]" box="[162,385,1078,1098]" pageId="4" pageNumber="5">
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<taxonomicName authority="Roksandic & Radović & Wu & Bae, 2021" authorityName="Roksandic & Radović & Wu & Bae" authorityYear="2021" box="[162,308,1078,1098]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="bodoensis" status="sp. nov.">
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<emphasis box="[162,308,1078,1098]" italics="true" pageId="4" pageNumber="5">Homo bodoensis</emphasis>
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</taxonomicName>
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<taxonomicNameLabel box="[313,385,1078,1098]" pageId="4" pageNumber="5" rank="species">sp. nov.</taxonomicNameLabel>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="4" pageNumber="5" type="etymology">
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<paragraph blockId="4.[122,771,1144,1965]" pageId="4" pageNumber="5">
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<emphasis box="[162,254,1144,1164]" italics="true" pageId="4" pageNumber="5">Etymology</emphasis>
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: The name
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<taxonomicName authorityName="Roksandic & Radović & Wu & Bae" authorityYear="2021" box="[373,460,1144,1164]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="bodoensis">
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<emphasis box="[373,460,1144,1164]" italics="true" pageId="4" pageNumber="5">bodoensis</emphasis>
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</taxonomicName>
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refers to the site of Bodo D'ar where the fossil specimen Bodo 1 was discovered.
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<bibRefCitation author="Conroy GC & Jolly CJ & Cramer D & Kalb JE" box="[586,613,1175,1188]" journalOrPublisher="Nature." pageId="4" pageNumber="5" pagination="67 - 70" part="276" refId="ref10946" refString="[123] Conroy GC, Jolly CJ, Cramer D, Kalb JE. Newly discovered fossil hominid skull from the Afar depression, Ethiopia. Nature. 1978; 276: 67 - 70." title="Newly discovered fossil hominid skull from the Afar depression, Ethiopia" type="journal article" year="1978">
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<superScript attach="left" box="[586,613,1175,1188]" fontSize="5" pageId="4" pageNumber="5">123</superScript>
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</bibRefCitation>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="4" pageNumber="5" type="materials_examined">
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<paragraph blockId="4.[122,771,1144,1965]" pageId="4" pageNumber="5">
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<materialsCitation ID-GBIF-Occurrence="3395527301" collectorName="Alemayehu Asfaw & Paul Whitehead" country="Ethiopia" location="Bodo D'ar" locationDeviation="Middle Awash research area, Afar Depression, the northwestern part of the former Hararghe Provice, Ethiopia" pageId="4" pageNumber="5" specimenCount="1" specimenCount-adult="1" specimenCount-male="1" typeStatus="holotype">
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<emphasis box="[162,242,1217,1237]" italics="true" pageId="4" pageNumber="5">
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<typeStatus box="[162,242,1217,1237]" pageId="4" pageNumber="5">Holotype</typeStatus>
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</emphasis>
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: Bodo 1, a partial cranium of an
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<specimenCount box="[549,595,1217,1237]" pageId="4" pageNumber="5" type="adult">adult</specimenCount>
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(presumably male) individual, preserving the face and the anterior braincase, found in autumn 1976 by
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<collectorName box="[284,454,1286,1306]" pageId="4" pageNumber="5">Alemayehu Asfaw</collectorName>
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,
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<collectorName box="[463,611,1286,1306]" pageId="4" pageNumber="5">Paul Whitehead</collectorName>
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and other members of the Rift Valley Research Mission in
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<collectingCountry box="[544,620,1321,1341]" name="Ethiopia" pageId="4" pageNumber="5">Ethiopia</collectingCountry>
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headed by Jon Kalb.
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<superScript attach="left" box="[168,226,1351,1364]" fontSize="5" pageId="4" pageNumber="5">123,124</superScript>
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The specimen is currently curated in the National Museum of Ethiopia in Addis Ababa, Ethiopia.
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<taxonomicName authorityName="Roksandic & Radović & Wu & Bae" authorityYear="2021" box="[563,678,1390,1410]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="bodoensis">
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<emphasis box="[563,578,1390,1410]" italics="true" pageId="4" pageNumber="5">H</emphasis>
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.
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<emphasis box="[591,678,1390,1410]" italics="true" pageId="4" pageNumber="5">bodoensis</emphasis>
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</taxonomicName>
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has been deposited in the ZooBank database (http://zoobank.org/) with Life Science Identifier urn:lsid:zoobank.org:act:
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<accessionNumber pageId="4" pageNumber="5">50AC3EA4-82E0-4AAD-BCDA-6DE6055888A7</accessionNumber>
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.
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</materialsCitation>
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</paragraph>
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</subSubSection>
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<caption ID-DOI="http://doi.org/10.5281/zenodo.5627847" ID-Zenodo-Dep="5627847" httpUri="https://zenodo.org/record/5627847/files/figure.png" pageId="4" pageNumber="5" startId="4.[818,830,1254,1273]" targetBox="[852,1434,141,1212]" targetPageId="4">
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<paragraph blockId="4.[818,1418,1254,1332]" pageId="4" pageNumber="5">
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<emphasis bold="true" box="[818,930,1254,1273]" pageId="4" pageNumber="5">FIGURE 1</emphasis>
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Asimplified model for the evolution of the genus
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<taxonomicName authorityName="Linnaeus" authorityYear="1758" box="[818,872,1283,1303]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
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<emphasis box="[818,872,1283,1303]" italics="true" pageId="4" pageNumber="5">Homo</emphasis>
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</taxonomicName>
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over the last 2 million years, with
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<taxonomicName authority="Roksandic & Radović & Wu & Bae, 2021" authorityName="Roksandic & Radović & Wu & Bae" authorityYear="2021" box="[1187,1333,1283,1303]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="bodoensis" status="sp. nov.">
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<emphasis box="[1187,1333,1283,1303]" italics="true" pageId="4" pageNumber="5">Homo bodoensis</emphasis>
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</taxonomicName>
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<taxonomicNameLabel box="[1339,1411,1283,1303]" pageId="4" pageNumber="5" rank="species">sp. nov.</taxonomicNameLabel>
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positioned as the ancestral (mostly African) form of
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<taxonomicName authorityName="Linnaeus" authorityYear="1758" box="[1294,1418,1312,1332]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="sapiens">
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<emphasis box="[1294,1418,1312,1332]" italics="true" pageId="4" pageNumber="5">Homo sapiens</emphasis>
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</taxonomicName>
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</paragraph>
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</caption>
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<subSubSection lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5" type="description">
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<paragraph blockId="4.[122,771,1144,1965]" lastBlockId="5.[122,771,1078,1999]" lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5">
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<emphasis box="[162,262,1529,1549]" italics="true" pageId="4" pageNumber="5">Description</emphasis>
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(modified from References 20,30,35,123,125): Bodo 1 comprises a damaged facial skeleton, partial neurocranium, and basicranium anterior to the basion of a single individual, reconstructed from dozens of individual bone fragments (
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<figureCitation box="[530,611,1633,1653]" captionStart="FIGURE" captionStartId="5.[122,134,892,911]" captionTargetBox="[124,768,138,851]" captionTargetId="figure-835@5.[124,768,138,851]" captionTargetPageId="5" captionText="FIGURE 2 Homo bodoensis sp. nov. holotype partial cranium Bodo 1 (Middle Awash, Ethiopia). Frontal (a), left lateral (b), superior (c) inferior (d) views. Scale bar: 5 cm. Source: Original photos Copyright © Jeffrey H. Schwartz" figureDoi="http://doi.org/10.5281/zenodo.5627849" httpUri="https://zenodo.org/record/5627849/files/figure.png" pageId="4" pageNumber="5">Figure 2</figureCitation>
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). Aside from the fact that the lateral portion of the right maxilla, the right zygomatic bone, and the left temporal process are missing, the face is generally well preserved. The palate is missing the portion posterior to the P4, and except for some small fragments of the right molar roots, the teeth are not preserved, and the alveolar processes show damage. The neurocranium preserves an almost complete frontal bone, the sphenoid, parts of the left temporal and both parietals, and the right portion of the occipital bone. The basicranial portion includes the partially preserved left mandibular fossa and articular eminence, the basioccipital, and the petrous portion of the temporal bone. The face is strikingly massive, with large rectangular orbits and a very broad interorbital region, a wide nasal root and aperture, a deep and robust left zygomatic, and a broad and deep palate. Though projecting and heavily built, the supraorbital tori are arched, segmented (i.e., divided into medial and lateral segments), and attenuated laterally; they do not form a continuous bony shelf but are rather separated by a prominent glabellar region, behind which is a flattened plane (rather than a sulcus). There is a distinct sagittal keeling in the frontal view, especially in the bregmatic region of the vault. The maxillary sinus is expanded, and there is no canine fossa. The frontal sinuses are also extensive and asymmetrical (the right sinus is larger). In lateral view, the skull is long and low, and the frontal presents a low and flattened profile. There is a prominent parietal angular torus, and the temporal squama is high and arched. The anterior nasal aperture is almost vertical in the lateral projection. In the superior view, the skull presents a piriform shape, broadening posteriorly from the noticeable postorbital constriction. In the inferior view, the large incisive foramen is placed anterior on the hard palate, the mandibular fossa is shallow, and the preserved part of the articular eminence is flat; the petrous portion of the temporal is placed in such a way that the foramen lacerum displays a crevice-like configuration. The endocranial capacity was estimated to ~1250 cm
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<bibRefCitation author="Prufer K & Racimo F & Patterson N" box="[318,327,1282,1295]" journalOrPublisher="Nature." pageId="5" pageNumber="6" pagination="43 - 49" part="505" refId="ref6772" refString="[3] Prufer K, Racimo F, Patterson N, et al. The complete genome sequence of a Neanderthal from the Altai Mountains. Nature. 2014; 505: 43 - 49." title="The complete genome sequence of a Neanderthal from the Altai Mountains" type="journal article" year="2014">
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<superScript attach="left" box="[318,327,1282,1295]" fontSize="5" pageId="5" pageNumber="6">3</superScript>
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</bibRefCitation>
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(i.e., between ~1200 and 1325 cm
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<bibRefCitation author="Prufer K & Racimo F & Patterson N" box="[677,686,1282,1295]" journalOrPublisher="Nature." pageId="5" pageNumber="6" pagination="43 - 49" part="505" refId="ref6772" refString="[3] Prufer K, Racimo F, Patterson N, et al. The complete genome sequence of a Neanderthal from the Altai Mountains. Nature. 2014; 505: 43 - 49." title="The complete genome sequence of a Neanderthal from the Altai Mountains" type="journal article" year="2014">
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<superScript attach="right" box="[677,686,1282,1295]" fontSize="5" pageId="5" pageNumber="6">3</superScript>
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</bibRefCitation>
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).
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<bibRefCitation author="Conroy GC & Weber GW & Seidler H & Recheis W & Zur Nedden D & Mariam JH" box="[696,723,1282,1295]" journalOrPublisher="Am J Phys Anthropol." pageId="5" pageNumber="6" pagination="111 - 118" part="113" refId="ref11010" refString="[125] Conroy GC, Weber GW, Seidler H, Recheis W, Zur Nedden D, Mariam JH. Endocranial capacity of the Bodo cranium determined from three-dimensional computed tomography. Am J Phys Anthropol. 2000; 113: 111 - 118." title="Endocranial capacity of the Bodo cranium determined from three-dimensional computed tomography" type="journal article" year="2000">
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<superScript attach="left" box="[696,723,1282,1295]" fontSize="5" pageId="5" pageNumber="6">125</superScript>
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</bibRefCitation>
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The series of cut marks situated on the facial and posterior parietal regions were interpreted as intentional postmortem defleshing.
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<bibRefCitation author="White TD" box="[630,657,1351,1364]" journalOrPublisher="Am J Phys Anthropol." pageId="5" pageNumber="6" pagination="503 - 509" part="69" refId="ref11057" refString="[126] White TD. Cut marks on the Bodo cranium: a case of prehistoric defleshing. Am J Phys Anthropol. 1986; 69: 503 - 509." title="Cut marks on the Bodo cranium: a case of prehistoric defleshing" type="journal article" year="1986">
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<superScript attach="left" box="[630,657,1351,1364]" fontSize="5" pageId="5" pageNumber="6">126</superScript>
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</bibRefCitation>
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</paragraph>
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</subSubSection>
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<caption ID-DOI="http://doi.org/10.5281/zenodo.5627849" ID-Zenodo-Dep="5627849" httpUri="https://zenodo.org/record/5627849/files/figure.png" pageId="5" pageNumber="6" startId="5.[122,134,892,911]" targetBox="[124,768,138,851]" targetPageId="5">
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<paragraph blockId="5.[122,745,892,999]" pageId="5" pageNumber="6">
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<emphasis bold="true" box="[122,233,892,911]" pageId="5" pageNumber="6">FIGURE 2</emphasis>
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<taxonomicName authority="Roksandic & Radović & Wu & Bae, 2021" authorityName="Roksandic & Radović & Wu & Bae" authorityYear="2021" box="[263,409,892,912]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="bodoensis" status="sp. nov.">
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<emphasis box="[263,409,892,912]" italics="true" pageId="5" pageNumber="6">Homo bodoensis</emphasis>
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</taxonomicName>
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<taxonomicNameLabel box="[414,487,892,912]" pageId="5" pageNumber="6" rank="species">sp. nov.</taxonomicNameLabel>
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holotype partial cranium Bodo 1 (Middle Awash, Ethiopia). Frontal (a), left lateral (b), superior (c) inferior (d) views. Scale bar: 5 cm.
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<emphasis box="[468,527,950,970]" italics="true" pageId="5" pageNumber="6">Source</emphasis>
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: Original photos Copyright © Jeffrey H. Schwartz
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</paragraph>
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</caption>
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<subSubSection pageId="5" pageNumber="6" type="materials_examined">
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<paragraph blockId="5.[122,771,1078,1999]" pageId="5" pageNumber="6">
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<emphasis box="[162,277,1390,1410]" italics="true" pageId="5" pageNumber="6">Type locality</emphasis>
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: Bodo D'ar, the Middle Awash research area, Afar Depression, the northwestern part of the former Hararghe Province, Ethiopia.
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</paragraph>
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<paragraph blockId="5.[122,771,1078,1999]" pageId="5" pageNumber="6">
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<emphasis box="[162,563,1494,1514]" italics="true" pageId="5" pageNumber="6">Geological age and stratigraphic position</emphasis>
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: Upper Bodo Sand Unit.
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<bibRefCitation author="Conroy GC & Jolly CJ & Cramer D & Kalb JE" box="[166,193,1524,1537]" journalOrPublisher="Nature." pageId="5" pageNumber="6" pagination="67 - 70" part="276" refId="ref10946" refString="[123] Conroy GC, Jolly CJ, Cramer D, Kalb JE. Newly discovered fossil hominid skull from the Afar depression, Ethiopia. Nature. 1978; 276: 67 - 70." title="Newly discovered fossil hominid skull from the Afar depression, Ethiopia" type="journal article" year="1978">
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<superScript attach="left" box="[166,193,1524,1537]" fontSize="5" pageId="5" pageNumber="6">123</superScript>
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</bibRefCitation>
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Dated to ca. 600 ka by laser-fusion
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<bibRefCitation author="Berger LR & Hawks J." box="[576,594,1524,1537]" journalOrPublisher="Am J Phys Anthropol." pageId="5" pageNumber="6" pagination="383 - 387" part="168" refId="ref7968" refString="[40] Berger LR, Hawks J. Australopithecus prometheus is a nomen nudum. Am J Phys Anthropol. 2019; 168: 383 - 387." title="Australopithecus prometheus is a nomen nudum" type="journal article" year="2019">
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<superScript attach="right" box="[576,594,1524,1537]" fontSize="5" pageId="5" pageNumber="6">40</superScript>
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</bibRefCitation>
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Ar/
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<superScript attach="right" box="[625,643,1524,1537]" fontSize="5" pageId="5" pageNumber="6">39</superScript>
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Ar technique (0.64 ± 0.03 Ma), biostratigraphy and tephrochronology.
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<bibRefCitation author="Clark JD & de Heinzelin J & Schick KD" box="[640,667,1559,1572]" journalOrPublisher="Science." pageId="5" pageNumber="6" pagination="1907 - 1910" part="264" refId="ref11089" refString="[127] Clark JD, de Heinzelin J, Schick KD, et al. African Homo erectus: old radiometric ages and young Oldowan assemblages in the middle Awash Valley, Ethiopia. Science. 1994; 264: 1907 - 1910." title="et al. African Homo erectus: old radiometric ages and young Oldowan assemblages in the middle Awash Valley, Ethiopia" type="journal article" year="1994">
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<superScript attach="left" box="[640,667,1559,1572]" fontSize="5" pageId="5" pageNumber="6">127</superScript>
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</bibRefCitation>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="5" pageNumber="6" type="discussion">
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<paragraph blockId="5.[122,771,1078,1999]" pageId="5" pageNumber="6">
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<emphasis box="[162,362,1598,1618]" italics="true" pageId="5" pageNumber="6">Archaeological context</emphasis>
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: The specimen is associated with an Acheulean stone tool assemblage.
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<superScript attach="left" box="[375,433,1628,1641]" fontSize="5" pageId="5" pageNumber="6">123,125</superScript>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="5" pageNumber="6" type="diagnosis">
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<paragraph blockId="5.[122,771,1078,1999]" lastBlockId="5.[818,1468,142,1791]" pageId="5" pageNumber="6">
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<emphasis box="[162,314,1667,1687]" italics="true" pageId="5" pageNumber="6">Species diagnosis</emphasis>
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: The species is diagnosed by a unique combination of cranial traits. The Bodo specimen has already been described as showing a mix of
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<taxonomicName authorityName="Dubois" authorityYear="1892" box="[309,398,1737,1757]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="erectus">
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<emphasis box="[309,324,1737,1757]" italics="true" pageId="5" pageNumber="6">H</emphasis>
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.
|
||
<emphasis box="[334,398,1737,1757]" italics="true" pageId="5" pageNumber="6">erectus</emphasis>
|
||
</taxonomicName>
|
||
-like and
|
||
<taxonomicName authorityName="Linnaeus" authorityYear="1758" box="[482,573,1737,1757]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="sapiens">
|
||
<emphasis box="[482,497,1737,1757]" italics="true" pageId="5" pageNumber="6">H</emphasis>
|
||
.
|
||
<emphasis box="[508,573,1737,1757]" italics="true" pageId="5" pageNumber="6">sapiens</emphasis>
|
||
</taxonomicName>
|
||
-like features.
|
||
<superScript attach="left" box="[698,769,1732,1745]" fontSize="5" pageId="5" pageNumber="6">
|
||
20,35,
|
||
<bibRefCitation author="Conroy GC & Jolly CJ & Cramer D & Kalb JE" box="[742,769,1732,1745]" journalOrPublisher="Nature." pageId="5" pageNumber="6" pagination="67 - 70" part="276" refId="ref10946" refString="[123] Conroy GC, Jolly CJ, Cramer D, Kalb JE. Newly discovered fossil hominid skull from the Afar depression, Ethiopia. Nature. 1978; 276: 67 - 70." title="Newly discovered fossil hominid skull from the Afar depression, Ethiopia" type="journal article" year="1978">123</bibRefCitation>
|
||
</superScript>
|
||
The species is similar to
|
||
<taxonomicName authorityName="Dubois" authorityYear="1892" box="[351,440,1771,1791]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="erectus">
|
||
<emphasis box="[351,366,1771,1791]" italics="true" pageId="5" pageNumber="6">H</emphasis>
|
||
.
|
||
<emphasis box="[377,440,1771,1791]" italics="true" pageId="5" pageNumber="6">erectus</emphasis>
|
||
</taxonomicName>
|
||
in having: a robustly built midface; total facial prognathism
|
||
<bibRefCitation author="Cartmill M & Smith FH" box="[341,368,1802,1815]" journalOrPublisher="John Wiley & Sons" pageId="5" pageNumber="6" refId="ref11134" refString="[128] Cartmill M, Smith FH. The Human Lineage. John Wiley & Sons; 2009." title="The Human Lineage" type="book" year="2009">
|
||
<superScript attach="left" box="[341,368,1802,1815]" fontSize="5" pageId="5" pageNumber="6">128</superScript>
|
||
</bibRefCitation>
|
||
; projecting tori and a flattened low frontal squama; sagittal keeling; a low vault profile; a prominent parietal angular torus; thick vault bones; no foramen lacerum is observable
|
||
<emphasis box="[740,757,1876,1895]" italics="true" pageId="5" pageNumber="6">—</emphasis>
|
||
it is presented as a narrow crevice.
|
||
<superScript attach="left" box="[437,486,1906,1919]" fontSize="5" pageId="5" pageNumber="6">20,128</superScript>
|
||
These traits can be linked to the retention of the general cranial structure from
|
||
<taxonomicName authorityName="Dubois" authorityYear="1892" box="[611,704,1945,1965]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="erectus">
|
||
<emphasis box="[611,626,1945,1965]" italics="true" pageId="5" pageNumber="6">H</emphasis>
|
||
.
|
||
<emphasis box="[640,704,1945,1965]" italics="true" pageId="5" pageNumber="6">erectus</emphasis>
|
||
</taxonomicName>
|
||
. Traits similar to other Middle Pleistocene and later hominin taxa include: increased cranial capacity and associated traits (broader frontal and mid-vault, reduced postorbital constriction, signs of parietal bossing, high and arched temporal squama), a vertical (rather than forward sloping) nasal margin, and the position of the incisive canal in front of the hard palate.
|
||
<superScript attach="left" box="[974,1046,276,289]" fontSize="5" pageId="5" pageNumber="6">
|
||
20,99,
|
||
<bibRefCitation author="Cartmill M & Smith FH" box="[1019,1046,276,289]" journalOrPublisher="John Wiley & Sons" pageId="5" pageNumber="6" refId="ref11134" refString="[128] Cartmill M, Smith FH. The Human Lineage. John Wiley & Sons; 2009." title="The Human Lineage" type="book" year="2009">128</bibRefCitation>
|
||
</superScript>
|
||
Excessively thick and projecting, but segmented brow ridges, with the incipient division of the brow at midorbit and attenuated laterally may be considered a distinctive trait of the species.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="5" pageNumber="6" type="discussion">
|
||
<paragraph blockId="5.[818,1468,142,1791]" pageId="5" pageNumber="6">
|
||
<emphasis box="[858,971,419,439]" italics="true" pageId="5" pageNumber="6">Comparisons</emphasis>
|
||
: In comparison to
|
||
<taxonomicName box="[1151,1233,419,439]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="erectu">
|
||
<emphasis box="[1151,1166,419,439]" italics="true" pageId="5" pageNumber="6">H</emphasis>
|
||
.
|
||
<emphasis box="[1178,1233,419,439]" italics="true" pageId="5" pageNumber="6">erectu</emphasis>
|
||
</taxonomicName>
|
||
s,
|
||
<taxonomicName authorityName="Roksandic & Radović & Wu & Bae" authorityYear="2021" box="[1255,1369,419,439]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="bodoensis">
|
||
<emphasis box="[1255,1270,419,439]" italics="true" pageId="5" pageNumber="6">H</emphasis>
|
||
.
|
||
<emphasis box="[1282,1369,419,439]" italics="true" pageId="5" pageNumber="6">bodoensis</emphasis>
|
||
</taxonomicName>
|
||
differs by the increased cranial capacity (intermediate between
|
||
<taxonomicName authorityName="Dubois" authorityYear="1892" box="[1311,1401,454,474]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="erectus">
|
||
<emphasis box="[1311,1326,454,474]" italics="true" pageId="5" pageNumber="6">H</emphasis>
|
||
.
|
||
<emphasis box="[1337,1401,454,474]" italics="true" pageId="5" pageNumber="6">erectus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Linnaeus" authorityYear="1758" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="sapiens">
|
||
<emphasis box="[1446,1461,454,474]" italics="true" pageId="5" pageNumber="6">H</emphasis>
|
||
.
|
||
<emphasis box="[818,883,489,509]" italics="true" pageId="5" pageNumber="6">sapiens</emphasis>
|
||
</taxonomicName>
|
||
) and a suite of associated derived traits: the curvature of the temporal squama; broader mid-vault; signs of parietal bossing; and relatively broad frontal bone where the maximal cranial breadth lies above the lower third of the skull in posterior view, with more vertical parietal walls.
|
||
</paragraph>
|
||
<paragraph blockId="5.[818,1468,142,1791]" pageId="5" pageNumber="6">
|
||
Increased cranial capacity is shared among most of the Middle Pleistocene hominins (excluding
|
||
<taxonomicName box="[1119,1198,697,717]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="naledi">
|
||
<emphasis box="[1119,1134,697,717]" italics="true" pageId="5" pageNumber="6">H</emphasis>
|
||
.
|
||
<emphasis box="[1145,1198,697,717]" italics="true" pageId="5" pageNumber="6">naledi</emphasis>
|
||
</taxonomicName>
|
||
and island isolates of Southeast Asia such as
|
||
<taxonomicName box="[981,1102,731,751]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="floresiensis">
|
||
<emphasis box="[981,996,731,751]" italics="true" pageId="5" pageNumber="6">H</emphasis>
|
||
.
|
||
<emphasis box="[1007,1102,731,751]" italics="true" pageId="5" pageNumber="6">floresiensis</emphasis>
|
||
</taxonomicName>
|
||
). This trait is presumably already under selection in the MRCA in the latter portion of the Early Pleistocene.
|
||
<superScript attach="left" box="[867,925,796,809]" fontSize="5" pageId="5" pageNumber="6">107,129</superScript>
|
||
Other features are not shared with Middle Pleistocene hominins such as
|
||
<taxonomicName authorityName="King" authorityYear="1864" box="[989,1165,835,855]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="neanderthalensis">
|
||
<emphasis box="[989,1004,835,855]" italics="true" pageId="5" pageNumber="6">H</emphasis>
|
||
.
|
||
<emphasis box="[1017,1165,835,855]" italics="true" pageId="5" pageNumber="6">neanderthalensis</emphasis>
|
||
</taxonomicName>
|
||
, late
|
||
<taxonomicName authorityName="Dubois" authorityYear="1892" box="[1221,1313,835,855]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="erectus">
|
||
<emphasis box="[1221,1236,835,855]" italics="true" pageId="5" pageNumber="6">H</emphasis>
|
||
.
|
||
<emphasis box="[1249,1313,835,855]" italics="true" pageId="5" pageNumber="6">erectus</emphasis>
|
||
</taxonomicName>
|
||
, and potentially other Asian groups yet to be systematized. The species differs from
|
||
<taxonomicName authorityName="King" authorityYear="1864" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="neanderthalensis">
|
||
<emphasis box="[1446,1461,870,890]" italics="true" pageId="5" pageNumber="6">H</emphasis>
|
||
.
|
||
<emphasis box="[818,966,905,925]" italics="true" pageId="5" pageNumber="6">neanderthalensis</emphasis>
|
||
</taxonomicName>
|
||
as it does not show any of the Neanderthal-specific morphology associated with midfacial prognathism and neurocranial shape. It also differs in the particular form of the brow ridges, which are smoothly continuous and double-arched in
|
||
<taxonomicName authorityName="King" authorityYear="1864" box="[1253,1426,1009,1029]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="neanderthalensis">
|
||
<emphasis box="[1253,1268,1009,1029]" italics="true" pageId="5" pageNumber="6">H</emphasis>
|
||
.
|
||
<emphasis box="[1278,1426,1009,1029]" italics="true" pageId="5" pageNumber="6">neanderthalensis</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
<bibRefCitation author="Schwartz JH & Tattersall I." box="[1431,1449,1004,1017]" journalOrPublisher="Am J Phys Anthropol." pageId="5" pageNumber="6" pagination="94 - 121" part="143" refId="ref7715" refString="[32] Schwartz JH, Tattersall I. Fossil evidence for the origin of Homo sapiens. Am J Phys Anthropol. 2010; 143: 94 - 121." title="Fossil evidence for the origin of Homo sapiens" type="journal article" year="2010">
|
||
<superScript attach="left" box="[1431,1449,1004,1017]" fontSize="5" pageId="5" pageNumber="6">32</superScript>
|
||
</bibRefCitation>
|
||
</paragraph>
|
||
<paragraph blockId="5.[818,1468,142,1791]" pageId="5" pageNumber="6">
|
||
<taxonomicName authorityName="Roksandic & Radović & Wu & Bae" authorityYear="2021" box="[858,973,1043,1063]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="bodoensis">
|
||
<emphasis box="[858,873,1043,1063]" italics="true" pageId="5" pageNumber="6">H</emphasis>
|
||
.
|
||
<emphasis box="[886,973,1043,1063]" italics="true" pageId="5" pageNumber="6">bodoensis</emphasis>
|
||
</taxonomicName>
|
||
lacks a number of the
|
||
<taxonomicName box="[1195,1279,1043,1063]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="sapien">
|
||
<emphasis box="[1195,1210,1043,1063]" italics="true" pageId="5" pageNumber="6">H</emphasis>
|
||
.
|
||
<emphasis box="[1223,1279,1043,1063]" italics="true" pageId="5" pageNumber="6">sapien</emphasis>
|
||
</taxonomicName>
|
||
s specific features
|
||
<emphasis box="[1449,1466,1044,1063]" italics="true" pageId="5" pageNumber="6">—</emphasis>
|
||
warranting a separate species designation. This is contrary to what is observed in
|
||
<taxonomicName authorityName="King" authorityYear="1864" box="[940,1119,1113,1133]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="neanderthalensis">
|
||
<emphasis box="[940,955,1113,1133]" italics="true" pageId="5" pageNumber="6">H</emphasis>
|
||
.
|
||
<emphasis box="[971,1119,1113,1133]" italics="true" pageId="5" pageNumber="6">neanderthalensis</emphasis>
|
||
</taxonomicName>
|
||
where the autapomorphies emerge early in the Middle Pleistocene. However, all of the later
|
||
<taxonomicName authorityName="Linnaeus" authorityYear="1758" box="[1372,1466,1147,1167]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="sapiens">
|
||
<emphasis box="[1372,1387,1147,1167]" italics="true" pageId="5" pageNumber="6">H</emphasis>
|
||
.
|
||
<emphasis box="[1401,1466,1147,1167]" italics="true" pageId="5" pageNumber="6">sapiens</emphasis>
|
||
</taxonomicName>
|
||
specific features can be derived from traits present in
|
||
<taxonomicName authorityName="Roksandic & Radović & Wu & Bae" authorityYear="2021" box="[1346,1461,1182,1202]" class="Mammalia" family="Hominidae" genus="Homo" kingdom="Animalia" order="Primates" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="bodoensis">
|
||
<emphasis box="[1346,1361,1182,1202]" italics="true" pageId="5" pageNumber="6">H</emphasis>
|
||
.
|
||
<emphasis box="[1374,1461,1182,1202]" italics="true" pageId="5" pageNumber="6">bodoensis</emphasis>
|
||
</taxonomicName>
|
||
, including the massive but segmented (divided into lateral and medial parts) browridges.
|
||
<superScript attach="left" box="[984,1024,1247,1260]" fontSize="5" pageId="5" pageNumber="6">20,34</superScript>
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="5" pageNumber="6" type="materials_examined">
|
||
<paragraph blockId="5.[818,1468,142,1791]" pageId="5" pageNumber="6">
|
||
<emphasis box="[858,950,1286,1306]" italics="true" pageId="5" pageNumber="6">Hypodigm</emphasis>
|
||
: In addition to the holotype Bodo 1, the hypodigm is based on the sufficiently preserved cranial specimens with the exclusion of isolated mandibles and includes
|
||
<emphasis box="[1257,1387,1355,1375]" italics="true" pageId="5" pageNumber="6">at a minimum</emphasis>
|
||
: Kabwe 1 (Broken Hill), Ndutu, Saldanha (Elandsfontein), Ngaloba (LH 18), and potentially Salé in Africa.
|
||
<superScript attach="left" box="[1098,1219,1420,1434]" fontSize="5" pageId="5" pageNumber="6">
|
||
30,108,
|
||
<bibRefCitation author="Clarke RJ" box="[1154,1181,1420,1433]" journalOrPublisher="Nature." pageId="5" pageNumber="6" pagination="485 - 487" part="262" refId="ref11194" refString="[130] Clarke RJ. New cranium of Homo erectus from Lake Ndutu, Tanzania. Nature. 1976; 262: 485 - 487." title="New cranium of Homo erectus from Lake Ndutu, Tanzania" type="journal article" year="1976">130</bibRefCitation>
|
||
<emphasis box="[1181,1190,1422,1434]" italics="true" pageId="5" pageNumber="6">–</emphasis>
|
||
<bibRefCitation author="Hublin J-J" box="[1191,1219,1420,1433]" editor="Barnham L & Robson- Brown K" journalOrPublisher="Western Academic and Specialist Press Limited" pageId="5" pageNumber="6" pagination="99 - 121" refId="ref11371" refString="[135] Hublin J-J. Northwestern African Middle Pleistocene hominids and their bearing on the emergence of Homo sapiens. In: Barnham L, Robson- Brown K, eds. Human Roots: Africa and Asia in the Middle Pleistocene. Western Academic and Specialist Press Limited; 2001: 99 - 121." title="Northwestern African Middle Pleistocene hominids and their bearing on the emergence of Homo sapiens" type="journal volume" volumeTitle="Human Roots: Africa and Asia in the Middle Pleistocene" year="2001">135</bibRefCitation>
|
||
</superScript>
|
||
Kabwe 1 could represent a late survivor of the taxon.
|
||
<bibRefCitation author="Grun R & Pike A & McDermott F" box="[1084,1112,1455,1468]" journalOrPublisher="Nature." pageId="5" pageNumber="6" pagination="372 - 375" part="580" refId="ref11427" refString="[136] Grun R, Pike A, McDermott F, et al. Dating the skull from Broken Hill, Zambia, and its position in human evolution. Nature. 2020; 580: 372 - 375." title="et al. Dating the skull from Broken Hill, Zambia, and its position in human evolution" type="journal article" year="2020">
|
||
<superScript attach="left" box="[1084,1112,1455,1468]" fontSize="5" pageId="5" pageNumber="6">136</superScript>
|
||
</bibRefCitation>
|
||
Some Middle Pleistocene specimens from Europe (e.g., Ceprano calvarium
|
||
<superScript attach="left" box="[1184,1244,1490,1503]" fontSize="5" pageId="5" pageNumber="6">137,138</superScript>
|
||
), could be included in this group as well. Locations, dating, previous taxonomic designations, and references for the included specimens are provided in the Table S1.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="5" pageNumber="6" type="distribution">
|
||
<paragraph blockId="5.[818,1468,142,1791]" pageId="5" pageNumber="6">
|
||
<emphasis box="[858,962,1633,1653]" italics="true" pageId="5" pageNumber="6">Distribution</emphasis>
|
||
: The species had a pan-African distribution with the peripheral range extending into the eastern Mediterranean (Southeast Europe and the Levant) from which it could have contributed to the repopulation of European (and possibly Central and East Asian) demographic sinks after the glaciations.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |