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<article article-type="brief-report" dtd-version="2.3" xml:lang="EN" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">
<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Earth Sci.</journal-id>
<journal-title>Frontiers in Earth Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Earth Sci.</abbrev-journal-title>
<issn pub-type="epub">2296-6463</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="publisher-id">1230707</article-id>
<article-id pub-id-type="doi">10.3389/feart.2023.1230707</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Earth Science</subject>
<subj-group>
<subject>Brief Research Report</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>A new assemblage of late Neanderthal remains from Cova Simanya (NE Iberia)</article-title>
<alt-title alt-title-type="left-running-head">Morales et al.</alt-title>
<alt-title alt-title-type="right-running-head">
<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/feart.2023.1230707">10.3389/feart.2023.1230707</ext-link>
</alt-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Morales</surname>
<given-names>Juan I.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<xref ref-type="corresp" rid="c001">&#x2a;</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2200181/overview"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Cebri&#xe0;</surname>
<given-names>Artur</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Soto</surname>
<given-names>Mar&#xed;a</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<xref ref-type="aff" rid="aff5">
<sup>5</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2345784/overview"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Rodr&#xed;guez-Hidalgo</surname>
<given-names>Antonio</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff6">
<sup>6</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2294397/overview"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Hernando</surname>
<given-names>Raquel</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2330151/overview"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Moreno-Ribas</surname>
<given-names>Elena</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2327897/overview"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Lombao</surname>
<given-names>Diego</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff7">
<sup>7</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1644114/overview"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Rabu&#xf1;al</surname>
<given-names>Jos&#x00E9; R.</given-names>
</name>
<xref ref-type="aff" rid="aff8">
<sup>8</sup>
</xref>
<xref ref-type="aff" rid="aff9">
<sup>9</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2330165/overview"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Mart&#xed;n-Perea</surname>
<given-names>David M.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff6">
<sup>6</sup>
</xref>
<xref ref-type="aff" rid="aff10">
<sup>10</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2354103/overview"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Garc&#x00ED;a-Tabernero</surname>
<given-names>Antonio</given-names>
</name>
<xref ref-type="aff" rid="aff11">
<sup>11</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2279239/overview"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Allu&#xe9;</surname>
<given-names>Ethel</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Garc&#x00ED;a-Basanta</surname>
<given-names>Andrea</given-names>
</name>
<xref ref-type="aff" rid="aff5">
<sup>5</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2328355/overview"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Lizano</surname>
<given-names>Esther</given-names>
</name>
<xref ref-type="aff" rid="aff12">
<sup>12</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/829131/overview"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Marqu&#x00E8;s-Bonet</surname>
<given-names>Tom&#x00E0;s</given-names>
</name>
<xref ref-type="aff" rid="aff12">
<sup>12</sup>
</xref>
<xref ref-type="aff" rid="aff13">
<sup>13</sup>
</xref>
<xref ref-type="aff" rid="aff14">
<sup>14</sup>
</xref>
<xref ref-type="aff" rid="aff15">
<sup>15</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Talamo</surname>
<given-names>Sahra</given-names>
</name>
<xref ref-type="aff" rid="aff16">
<sup>16</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/638593/overview"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Tassoni</surname>
<given-names>Laura</given-names>
</name>
<xref ref-type="aff" rid="aff16">
<sup>16</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Lalueza-Fox</surname>
<given-names>Carles</given-names>
</name>
<xref ref-type="aff" rid="aff13">
<sup>13</sup>
</xref>
<xref ref-type="aff" rid="aff17">
<sup>17</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1584129/overview"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Fullola</surname>
<given-names>Josep M.</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2366842/overview"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Rosas</surname>
<given-names>Antonio</given-names>
</name>
<xref ref-type="aff" rid="aff11">
<sup>11</sup>
</xref>
<xref ref-type="corresp" rid="c001">&#x2a;</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2351685/overview"/>
</contrib>
</contrib-group>
<aff id="aff1">
<sup>1</sup>
<institution>IPHES-CERCA</institution>, <institution>Institut Catal&#xe0; de Paleoecologia Humana i Evoluci&#xf3; Social</institution>, <addr-line>Tarragona</addr-line>, <country>Spain</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>Departament d&#x2019;Hist&#xf2;ria i Hist&#xf2;ria de l&#x2019;Art</institution>, <institution>Universitat Rovira i Virgili</institution>, <addr-line>Tarragona</addr-line>, <country>Spain</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>SERP</institution>, <institution>Departament d&#x27;Hist&#xf2;ria i Arqueologia</institution>, <institution>Secci&#xf3; de Prehistoria</institution>, <institution>Universitat de Barcelona</institution>, <addr-line>Barcelona</addr-line>, <country>Spain</country>
</aff>
<aff id="aff4">
<sup>4</sup>
<institution>Madrid Institute for Advanced Study, MIAS</institution>, <addr-line>Madrid</addr-line>, <country>Spain</country>
</aff>
<aff id="aff5">
<sup>5</sup>
<institution>Departamento de Prehistoria y Arqueolog&#xed;a</institution>, <institution>Facultad de Filosof&#xed;a y Letras</institution>, <institution>Universidad Aut&#xf3;noma de Madrid</institution>, <addr-line>Madrid</addr-line>, <country>Spain</country>
</aff>
<aff id="aff6">
<sup>6</sup>
<institution>Instituto de Evoluci&#xf3;n en &#xc1;frica (IDEA)</institution>, <addr-line>Madrid</addr-line>, <country>Spain</country>
</aff>
<aff id="aff7">
<sup>7</sup>
<institution>GEPN-AAT</institution>, <institution>Departamento de Historia</institution>, <institution>Facultade de Xeograf&#xed;a e Historia</institution>, <institution>Universidade Santiago de Compostela</institution>, <addr-line>Santiago de Compostela</addr-line>, <country>Spain</country>
</aff>
<aff id="aff8">
<sup>8</sup>
<institution>Instituto Universitario de Investigaci&#xf3;n en Arqueolog&#xed;a y Patrimonio Hist&#xf3;rico</institution>, <institution>Universidad de Alicante</institution>, <addr-line>Alicante</addr-line>, <country>Spain</country>
</aff>
<aff id="aff9">
<sup>9</sup>
<institution>Department of Archaeology and Heritage Studies</institution>, <institution>Aarhus University</institution>, <addr-line>H&#xf8;jbjerg</addr-line>, <country>Denmark</country>
</aff>
<aff id="aff10">
<sup>10</sup>
<institution>Department of Geodynamics, Stratigraphy and Palaeontology</institution>, <institution>Complutense University of Madrid</institution>, <addr-line>Madrid</addr-line>, <country>Spain</country>
</aff>
<aff id="aff11">
<sup>11</sup>
<institution>Group of Paleoanthropology</institution>, <institution>Museo Nacional de Ciencias Naturales (CSIC)</institution>, <addr-line>Madrid</addr-line>, <country>Spain</country>
</aff>
<aff id="aff12">
<sup>12</sup>
<institution>Institut Catal&#xe0; de Paleontologia Miquel Crusafont</institution>, <institution>Universitat Aut&#xf2;noma de Barcelona</institution>, <addr-line>Barcelona</addr-line>, <country>Spain</country>
</aff>
<aff id="aff13">
<sup>13</sup>
<institution>Institute of Evolutionary Biology (UPF-CSIC)</institution>, <institution>University Pompeu Fabra</institution>, <addr-line>Barcelona</addr-line>, <country>Spain</country>
</aff>
<aff id="aff14">
<sup>14</sup>
<institution>Catalan Institution of Research and Advanced Studies (ICREA)</institution>, <addr-line>Barcelona</addr-line>, <country>Spain</country>
</aff>
<aff id="aff15">
<sup>15</sup>
<institution>CNAG-CRG</institution>, <institution>Centre for Genomic Regulation (CRG)</institution>, <institution>Barcelona Institute of Science and Technology (BIST)</institution>, <addr-line>Barcelona</addr-line>, <country>Spain</country>
</aff>
<aff id="aff16">
<sup>16</sup>
<institution>Department of Chemistry Giacomo Ciamician</institution>, <institution>University of Bologna</institution>, <addr-line>Bologna</addr-line>, <country>Italy</country>
</aff>
<aff id="aff17">
<sup>17</sup>
<institution>Museum of Natural Sciences of Barcelona</institution>, <addr-line>Barcelona</addr-line>, <country>Spain</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>
<bold>Edited by:</bold> <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/551276/overview">Raymond Louis Bernor</ext-link>, Howard University, United States</p>
</fn>
<fn fn-type="edited-by">
<p>
<bold>Reviewed by:</bold> <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/1403114/overview">Leslea J. Hlusko</ext-link>, National Center for Research on Human Evolution (CENIEH), Spain</p>
<p>
<ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/1500082/overview">John De Vos</ext-link>, Naturalis Biodiversity Center, Netherlands</p>
<p>
<ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/497795/overview">Marina Melchionna</ext-link>, University of Naples Federico II, Italy</p>
</fn>
<corresp id="c001">&#x2a;Correspondence: Juan I. Morales, <email>jmorales@iphes.cat</email>; Antonio Rosas, <email>arosas@mncn.csic.es</email>
</corresp>
</author-notes>
<pub-date pub-type="epub">
<day>19</day>
<month>09</month>
<year>2023</year>
</pub-date>
<pub-date pub-type="collection">
<year>2023</year>
</pub-date>
<volume>11</volume>
<elocation-id>1230707</elocation-id>
<history>
<date date-type="received">
<day>29</day>
<month>05</month>
<year>2023</year>
</date>
<date date-type="accepted">
<day>04</day>
<month>07</month>
<year>2023</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2023 Morales, Cebri&#xe0;, Soto, Rodr&#xed;guez-Hidalgo, Hernando, Moreno-Ribas, Lombao, Rabu&#xf1;al, Mart&#xed;n-Perea, Garc&#x00ED;a-Tabernero, Allu&#xe9;, Garc&#x00ED;a-Basanta, Lizano, Marqu&#x00E8;s-Bonet, Talamo, Tassoni, Lalueza-Fox, Fullola and Rosas.</copyright-statement>
<copyright-year>2023</copyright-year>
<copyright-holder>Morales, Cebri&#xe0;, Soto, Rodr&#xed;guez-Hidalgo, Hernando, Moreno-Ribas, Lombao, Rabu&#xf1;al, Mart&#xed;n-Perea, Garc&#x00ED;a-Tabernero, Allu&#xe9;, Garc&#x00ED;a-Basanta, Lizano, Marqu&#x00E8;s-Bonet, Talamo, Tassoni, Lalueza-Fox, Fullola and Rosas</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>This study presents an exceptional collection of 54 Late Pleistocene human remains that correspond to at least three Neanderthal individuals from Simanya Gran, the main gallery of Cova Simanya, located in the northeastern Iberian Peninsula. The collection comprised 53 unpublished remains that were unearthed during the 1970s and an additional tooth discovered during 2021 excavations. The specimens represent an adult with a small stature, a periadolescent aged approximately 11.5&#xa0;years, and an immature individual aged approximately 7.7&#xa0;years, thus offering a more complete demographic perspective. The collection encompasses diverse anatomical parts including upper and lower dentition, mandible, vertebrae, and limb bones from both the upper and lower extremities. Attempts to extract aDNA were unsuccessful. Renewed archaeological investigations at Cova Simanya have facilitated the reevaluation of the original stratigraphic context of these remains, leading to the discovery of the additional tooth, aligning with the periadolescent individual. This assemblage is currently the most extensive Neanderthal collection from the northeastern Mediterranean Iberia, offering invaluable insights into the morphology and evolutionary trajectory of Late Pleistocene hominins. Hence, Simanya Neanderthals will enhance our understanding of Neanderthal demographics and evolution, paving the way for an in-depth examination of the morphological diversity and evolutionary context of Iberian Neanderthals.</p>
</abstract>
<kwd-group>
<kwd>Late Pleistocene</kwd>
<kwd>Middle Paleolithic archeology</kwd>
<kwd>human remains</kwd>
<kwd>Neanderthal</kwd>
<kwd>Cova Simanya</kwd>
<kwd>paleoanthropology</kwd>
<kwd>Paleolithic archaeology</kwd>
<kwd>Iberian Peninsula</kwd>
</kwd-group>
<contract-num rid="cn001">PID2021-122356NB-I00 PID2020-113960GB-I00 PID2021-124590NB-I00 PID2019-103987GB-C31 PID2021-126004NB-100 CEX2019-000945-M</contract-num>
<contract-num rid="cn002">2021 SGR 01237 2021 SGR 00337 2021 SGR 00177</contract-num>
<contract-num rid="cn003">Anal&#xed;ticas 2022-2023</contract-num>
<contract-num rid="cn004">803147 RESOLUTION 864203</contract-num>
<contract-num rid="cn005">ARQ001SOL-172-2022</contract-num>
<contract-num rid="cn006">ED481B-2022-048</contract-num>
<contract-sponsor id="cn001">Ministerio de Ciencia e Innovaci&#xf3;n<named-content content-type="fundref-id">10.13039/501100004837</named-content>
</contract-sponsor>
<contract-sponsor id="cn002">Ag&#xe8;ncia de Gesti&#xf3; d&#x27;Ajuts Universitaris i de Recerca<named-content content-type="fundref-id">10.13039/501100003030</named-content>
</contract-sponsor>
<contract-sponsor id="cn003">Fundaci&#xf3;n Palarq<named-content content-type="fundref-id">10.13039/501100015681</named-content>
</contract-sponsor>
<contract-sponsor id="cn004">HORIZON EUROPE European Research Council<named-content content-type="fundref-id">10.13039/100019180</named-content>
</contract-sponsor>
<contract-sponsor id="cn005">Generalitat de Catalunya<named-content content-type="fundref-id">10.13039/501100002809</named-content>
</contract-sponsor>
<contract-sponsor id="cn006">Xunta de Galicia<named-content content-type="fundref-id">10.13039/501100010801</named-content>
</contract-sponsor>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Paleontology</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1">
<title>1 Introduction</title>
<p>Neanderthals exhibited distinct genetic histories and cultural adaptations along time and space. In the Iberian Peninsula, the Iberian Mediterranean region holds significant importance in understanding Neanderthal evolution and variability, and its archaeological record provides valuable insights into their biological and cultural development as well as their eventual disappearance during marine isotope stage (MIS) 3.</p>
<p>Middle Paleolithic Mousterian sites can be found along the Mediterranean coast from Gibraltar to the Pyrenees, with Neanderthal remains being documented at several locations (<xref ref-type="bibr" rid="B16">Dean et al., 1986</xref>; <xref ref-type="bibr" rid="B3">Arsuaga and Berm&#xfa;dez de Castro, 1987</xref>; <xref ref-type="bibr" rid="B14">Daura et al., 2005</xref>; <xref ref-type="bibr" rid="B5">Arsuaga et al., 2007</xref>; <xref ref-type="bibr" rid="B44">Walker et al., 2008</xref>; <xref ref-type="bibr" rid="B13">Daura et al., 2010</xref>; <xref ref-type="bibr" rid="B45">Walker et al., 2010</xref>; <xref ref-type="bibr" rid="B31">Rodr&#xed;guez et al., 2011</xref>; <xref ref-type="bibr" rid="B4">Arsuaga et al., 2012</xref>; <xref ref-type="bibr" rid="B20">Maroto et al., 2012</xref>; <xref ref-type="bibr" rid="B23">Michel et al., 2013</xref>; <xref ref-type="bibr" rid="B2">Aparicio et al., 2014</xref>; <xref ref-type="bibr" rid="B17">Garralda et al., 2014</xref>; <xref ref-type="bibr" rid="B29">Quam et al., 2015</xref>; <xref ref-type="bibr" rid="B8">Bokelmann et al., 2019</xref>; <xref ref-type="bibr" rid="B11">Carri&#xf3;n et al., 2019</xref>) (<xref ref-type="fig" rid="F1">Figure 1A</xref>).</p>
<fig id="F1" position="float">
<label>FIGURE 1</label>
<caption>
<p>
<bold>(A)</bold> Map with the distribution of Neanderthal remains in the Iberian Peninsula; <bold>(B)</bold> main Middle Paleolithic sites from NE Iberia and the situation of Cova Simanya; <bold>(C)</bold> Cova Simanya karstic system, in red the areas excavated between 2020&#x2013;2022, and <bold>(D)</bold> main stratigraphic sections for the Pleistocene sequence.</p>
</caption>
<graphic xlink:href="feart-11-1230707-g001.tif"/>
</fig>
<p>These data suggest that Mediterranean Iberia was continuously inhabited by Neanderthals from MIS-7 onwards, with northeastern Iberia being an especially dense Mousterian cluster (<xref ref-type="bibr" rid="B21">Mart&#xed;nez-Moreno et al., 2010a</xref>; <xref ref-type="bibr" rid="B22">Mart&#xed;nez-Moreno et al., 2010b</xref>; <xref ref-type="bibr" rid="B36">Rosell et al., 2010</xref>; <xref ref-type="bibr" rid="B25">Mora et al., 2018</xref>; <xref ref-type="bibr" rid="B27">Morales et al., 2022</xref>) (<xref ref-type="fig" rid="F1">Figure 1B</xref>), including long-term occupation archives such as Abric Roman&#xed; (Barcelona) (<xref ref-type="bibr" rid="B7">Bischoff et al., 1988</xref>; <xref ref-type="bibr" rid="B42">Vallverd&#xfa; et al., 2010</xref>; <xref ref-type="bibr" rid="B10">Carbonell, 2012</xref>) or L&#x2019;Arbreda (<xref ref-type="bibr" rid="B39">Soler et al., 2014</xref>), and the southernmost occurrence of Ch&#xe2;telperronian technologies (<xref ref-type="bibr" rid="B26">Morales et al., 2019</xref>; <xref ref-type="bibr" rid="B32">Rodr&#xed;guez-Hidalgo et al., 2019</xref>). However, the paleoanthropological record in this region is restricted to the isolated teeth from Mollet (&#x223c;215&#xa0;ka) (<xref ref-type="bibr" rid="B20">Maroto et al., 2012</xref>), and the Cova del Gegant assemblage dated back to 55,000&#x2013;52,000&#xa0;years ago (<xref ref-type="bibr" rid="B14">Daura et al., 2005</xref>; <xref ref-type="bibr" rid="B13">Daura et al., 2010</xref>; <xref ref-type="bibr" rid="B29">Quam et al., 2015</xref>; <xref ref-type="bibr" rid="B15">Daura et al., 2022</xref>) once the long-time disputed Banyoles mandible does not show any derived Neanderthal features (<xref ref-type="bibr" rid="B19">Keeling et al., 2023</xref>).</p>
<p>Herein, we present a novel Neanderthal assemblage from Cova Simanya (Sant Lloren&#xe7; Savall, Barcelona), first recognized during the analysis of a museum collection and constituting the largest sample documented so far in northeastern Iberia. This discovery has led to a new research project that aimed at reconstructing the genesis, context, and paleobiology of the Neanderthal sample, including renewed excavations at Cova Simanya. These works have provided a robust archeological context for the uncontextualized &#x201c;Museum Collection&#x201d; including additional Neanderthal remains.</p>
<p>Simanya&#x2019;s assemblage may offer insights into the late Neanderthals&#x2019; apomorphic traits and their congruence with other known samples, while also raising questions about the genetic diversity and population turnovers within the Iberian Peninsula (<xref ref-type="bibr" rid="B43">Vernot et al., 2021</xref>).</p>
</sec>
<sec sec-type="materials|methods" id="s2">
<title>2 Materials and methods</title>
<sec id="s2-1">
<title>2.1 Cova Simanya</title>
<p>Cova Simanya is a karstic system (UTM ETRS89 417517&#x2013;4614066, 885 masl) opened in Montcau mountain, running through the Paleogene conglomerates forming the Sant Lloren&#xe7; del Munt Massif, on the eastern margin of the Ebro Basin, and has over 300&#xa0;m of graticulated galleries. The Font del Llor stream cuts the system, creating three consecutive entrances known as Simanya Triangle, Simanya Petita, and Simanya Gran (<xref ref-type="fig" rid="F1">Figure 1C</xref> and <xref ref-type="sec" rid="s10">Supplementary Figure S1</xref>).</p>
<p>Simanya Gran is the larger sector of Cova Simanya, comprising the Western Gallery (WG), Main Gallery (MG), Pas de l&#x2019;Oca Gallery (POG), and the inner part of the Main Gallery (IMG) (<xref ref-type="fig" rid="F1">Figure 1C</xref>). The cave has been known to scholars at least since the 18th century, and the presence of archeological remains is known since the first excavation in the 1930s.</p>
</sec>
<sec id="s2-2">
<title>2.2 The &#x201c;Museum Collection&#x201d;</title>
<p>The &#x201c;Museum Collection&#x201d; was collected by an amateur, Mr. Miguel Aznar, in 1978&#x2013;79 and contains a combination of two distinct sources: 1) surface findings and 2) remains from a small excavation. The surface findings include isolated human and animal bones, pottery, and chert fragments. The excavation yielded human and faunal remains, as well as a few samples of charcoal, sediment, and carbonate crusts. The excavation was performed in a breccia deposit of reddish sediments with abundant charcoals located in the IMG sector. No pottery or any other Holocene remains were found during the excavation. The collection containing 143 osteological remains was donated to the Museum of Archaeology of Catalonia in 1986. In 2021, Mr. Aznar delivered two additional bags to us containing small and mostly undiagnostic bone fragments that are not included in the original donation.</p>
<p>Despite being layered in the reddish clay-like sediment, the human bone collection exhibited a notable state of preservation. However, the presence of modern fractures and adhesive-induced alterations from the 1970s necessitated a thorough evaluation of their conservation status. Following the evaluation, a series of measures were taken to inhibit further deterioration. Cleaning procedures involving water and soft brushes were applied, and outdated adhesives were removed and substituted with Paraloid<sup>&#xae;</sup> B-72, a durable and reversible acrylic resin. This resin was also utilized to mend modern fractures and consolidate the overall structure, ensuring the assemblage&#x2019;s longevity and integrity for future study and preservation.</p>
</sec>
<sec id="s2-3">
<title>2.3 2020&#x2013;2022 excavations</title>
<p>The 2020&#x2013;2022 excavation programs targeted Simanya Gran and aimed at identifying the original context of the &#x201c;Museum Collection&#x201d; and evaluating the potential of the Pleistocene deposits in the cave. In 2020, five test pits were excavated to reconstruct the longitudinal stratigraphic development along the MG (S1, S2, and S3) and WG (S4&#x2013;S5). In 2021, a more extensive excavation was carried out in the IMG sector (<xref ref-type="fig" rid="F1">Figure 1D</xref>). The latter provided a similar fossil assemblage to the &#x201c;Museum Collection,&#x201d; including a new Neanderthal tooth (SI-60) found in the area reported by Mr. Aznar as the origin of the collection (<xref ref-type="sec" rid="s10">Supplementary Table S1</xref>). In 2022, we continued with the excavation of the IMG and expanded the S3 test pit of the MG to a surface of 10&#xa0;m<sup>2</sup>. A visual recognition of Mr. Aznar excavations (trenches, sections, or backfills) was not possible due to the homogenization of the sedimentary surface by the unrestricted public access.</p>
</sec>
<sec id="s2-4">
<title>2.4 Radiocarbon dating</title>
<p>Eleven bone samples, both anthropized and non-anthropized remains, and five charcoals (<italic>Pinus sylvestris</italic> type) from the &#x201c;Museum Collection&#x201d; and our own excavations at the IMG were selected for radiocarbon pretreatment at the University of Bologna&#x2019;s BRAVHO facilities. <xref ref-type="bibr" rid="B40">Talamo et al. (2021)</xref> followed the collagen extraction method, and only a non-anthropized <italic>Capra pyrenaica</italic> humerus (SG21.06.39) from the excavation yielded sufficient collagen for <sup>14</sup>C dating. Charcoal pretreatment used the ABOx method proposed by <xref ref-type="bibr" rid="B41">Tassoni et al. (2022)</xref>, adapting <xref ref-type="bibr" rid="B6">Bird et al.&#x2019;s (1999)</xref> method to small, poorly preserved samples by adjusting the HCl solution&#x2019;s molarity and reducing the oxidation step&#x2019;s reaction time. After the pretreatment, the collagen and charcoal samples were graphitized at the BRAVHO lab (<xref ref-type="bibr" rid="B41">Tassoni et al., 2022</xref>). During the charcoal sample SG-CA-200 combustion, the C integral peak obtained was very low for analysis and was excluded. The resulting graphite was pressed into targets and sent to the Mannheim AMS Laboratory at the Curt-Engelhorn-Center for Archeometry for measurement.</p>
<p>All <sup>14</sup>C ages were calibrated with the IntCal20 calibration curve (<xref ref-type="bibr" rid="B30">Reimer et al., 2020</xref>) using OxCal v4.4 software (<xref ref-type="bibr" rid="B9">Bronk Ramsey, 2009</xref>).</p>
</sec>
<sec id="s2-5">
<title>2.5 aDNA</title>
<p>An attempt to extract aDNA from Simanya Neanderthals was made by sampling a tooth and a phalanx from the &#x201c;Museum Collection&#x201d; through shotgun sequencing (Illumina NovaSeq 6000) and mitochondrial DNA (mtDNA) capture with Neanderthal-specific baits (Arbor Biosciences).</p>
</sec>
</sec>
<sec sec-type="results" id="s3">
<title>3 Results</title>
<sec id="s3-1">
<title>3.1 Stratigraphic context and archeo-paleontological record</title>
<p>The litho-stratigraphic sequence documented so far for the MG and IMG sectors of Simanya Gran contains a primary Pleistocene sequence capped by a thin Holocene deposit, 5&#x2013;30&#xa0;cm thick, which is documented from the entrance up to 53 meter. From that point of the MG to the end of the IMG, the Pleistocene units are uncapped (<xref ref-type="sec" rid="s10">Supplementary Figure S2.1</xref>) and four main litho-stratigraphic units have been described (<xref ref-type="fig" rid="F1">Figure 1D</xref>), namely, from the bottom to the top, Unit SG4 (cave clays), Unit SG3 (channel deposits), Unit SG2 (complex unit), and Unit SG1 (diamicton).</p>
<p>Unit SG4 contains at least 60&#xa0;cm of pure, dark reddish-brown clays with scarce coarse to very coarse gravels, mainly limestone and slate, documented at S1, S2, S3, and the IMG, and is eroded by Unit SG3. Unit SG3 comprised several coarsening upward sequences (silt/clay to coarse sand) that are cut and filled by at least four poorly sorted (silts to coarse sand) channel deposits. The thickness of Unit SG3 varies from 5 to 45&#xa0;cm, and it has been documented in S1, S2, S3, and the IMG. In the S1 section of the MG sector, Unit SG3 is conformably overlain by Unit SG2-a, an approximately 50&#xa0;cm-thick matrix-supported conglomerate of slate and limestone clasts that are more abundant toward the top, while in the S3 section, it is overlain by Units SG2.3-to-1, a succession of brown clays, a poorly sorted yellowish sandy deposit, and a very fine gravel conglomerate, and by the S1 flowstone. The Pleistocene sequence ends with Unit SG1, a reddish-brown, matrix-supported polygenic limestone, sandstone, and slate conglomerate with an erosive base that has been documented exclusively at the IMG.</p>
<p>Three archaeological layers have been documented so far within the litho-stratigraphic sequence at the MG&#x2013;IMG sector. Layer P1 corresponds to the Holocene phase and yields artifacts from the Late Neolithic to the Late Antiquity, including scattered funerary evidence dated back to the 5th century AD, in a condensed sub-superficial sedimentary palimpsest (<xref ref-type="sec" rid="s10">Supplementary Figure S2.1</xref>). The Pleistocene layer O300-PO400 (Unit SG1) was documented in the IMG and is characterized by a paleontological background composed mostly of <italic>Ursus</italic> cf<italic>. arctos</italic>, <italic>U</italic>. cf. <italic>spelaeus</italic>, and <italic>Capra pyrenaica</italic>. The preliminary analysis of the assemblage suggests a combination of taphonomic agents and processes in the formation of the record, including bear deaths during hibernation, accumulation of Iberian ibex by a large carnivore (likely a felid), and transportation and/or reworking by physical processes such as water flow and gravity. This taxonomic profile is consistent with the &#x201c;Museum Collection,&#x201d; which mostly consists of <italic>U</italic>. cf. <italic>arctos</italic> and <italic>Capra pyrenaica</italic>. Evidence of occasional Middle Paleolithic Mousterian occupation has also been found to be interspersed in the paleontological background of O300-PO400, over 70&#xa0;m inside the cave (<xref ref-type="sec" rid="s10">Supplementary Figures S2.2, 2.3, S3</xref>). The evidence associated to human occupations contains a small lithic assemblage and some faunal remains (<italic>Sus scrofa</italic> and <italic>Cervus elaphus</italic>) that exhibit cut marks or green breakage patterns. Within this horizon, a small and flat combustion feature was also documented. The presence of core-trimming flakes, a sidescraper, multifaceted butts, dorso-ventral right angles, and centripetal dorsal scar patterns in the lithic assemblage clearly indicates a Mousterian affinity (<xref ref-type="bibr" rid="B27">Morales et al., 2022</xref>). Additionally, a third upper human molar (SI-60) was found in Layer O300-PO400 (301&#x2013;80 meter). Finally, Layer P300, documented exclusively at S3, has provided a faunal assemblage composed of highly cemented skeletal remains of <italic>Capra pyrenaica</italic> and isolated remains of <italic>Cervus elaphus</italic>, without any evidence of human activity so far.</p>
</sec>
<sec id="s3-2">
<title>3.2 Dating</title>
<p>Calibrated dates at 68.3% and 95.4% probability are provided in <xref ref-type="table" rid="T1">Table 1</xref>. The bone sample is the only one that results in a final calibrated range of 42,440&#x2013;42,110&#xa0;cal BP (68.3%). All the other four charcoal samples are older than the radiocarbon method (&#x3e;49,000).</p>
<table-wrap id="T1" position="float">
<label>TABLE 1</label>
<caption>
<p>Samples selected for <sup>14</sup>C dating pretreated at the University of Bologna and AMS results.</p>
</caption>
<table>
<thead valign="top">
<tr>
<th align="center">Bologna Code</th>
<th align="center">Sample ID</th>
<th align="center">Unit</th>
<th align="center">Sector</th>
<th align="center">Material</th>
<th align="center">Species</th>
<th align="center">Collagen (mg)</th>
<th align="center">Collagen (%)</th>
<th align="center">AMS Code</th>
<th align="center">d13C</th>
<th align="center">F14C</th>
<th align="center">C:N</th>
<th align="center">14C age</th>
<th align="center">1s Error</th>
<th align="center">Cal BP age (68.3%)</th>
<th align="center">Cal BP age (95.4%)</th>
</tr>
</thead>
<tbody valign="top">
<tr>
<td align="center">BRA-6108</td>
<td align="center">SG21.06.39</td>
<td align="center">O300-PO400</td>
<td align="left">301-70</td>
<td align="center">Bone</td>
<td align="center">Caprini</td>
<td align="center">4.8</td>
<td align="center">1%</td>
<td align="center">MAMS 56486</td>
<td align="center">&#x2212;18.21</td>
<td align="center">0.0088</td>
<td align="center">2.89</td>
<td align="center">38009</td>
<td align="center">410</td>
<td align="left">42440&#x2013;42110</td>
<td align="left">42620&#x2013;41930</td>
</tr>
<tr>
<td align="center">BRA-6109</td>
<td align="center">SG21.11.08</td>
<td align="center">O300-PO400</td>
<td align="left">301-71</td>
<td align="center">Bone</td>
<td align="center">Suidae</td>
<td align="center">0.7</td>
<td align="center">0%</td>
<td align="center">&#x2014;</td>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
</tr>
<tr>
<td align="center">BRA-6110</td>
<td align="center">SG21.06.78</td>
<td align="center">O300-PO400</td>
<td align="left">301-82</td>
<td align="center">Bone</td>
<td align="center">Caprini</td>
<td align="center">0.6</td>
<td align="center">0%</td>
<td align="center">&#x2014;</td>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
</tr>
<tr>
<td align="center">BRA-6111</td>
<td align="center">SG21.10.58</td>
<td align="center">O300-PO400</td>
<td align="left">301-82</td>
<td align="center">Bone</td>
<td align="center">Caprini</td>
<td align="center">0.2</td>
<td align="center">0%</td>
<td align="center">&#x2014;</td>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
</tr>
<tr>
<td align="center">BRA-6112</td>
<td align="center">SG21.12.10</td>
<td align="center">O300-PO400</td>
<td align="left">301-82</td>
<td align="center">Tooth</td>
<td align="center">Caprini</td>
<td align="center">&#x2014;</td>
<td align="center">&#x2014;</td>
<td align="center">&#x2014;</td>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
</tr>
<tr>
<td align="center">BRA-6113</td>
<td align="center">SG21.10.30</td>
<td align="center">O300-PO400</td>
<td align="left">301-83</td>
<td align="center">Tooth</td>
<td align="center">Caprini</td>
<td align="center">&#x2014;</td>
<td align="center">&#x2014;</td>
<td align="center">&#x2014;</td>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
</tr>
<tr>
<td align="center">BRA-6114</td>
<td align="center">121_1978</td>
<td align="center">Museum Collection</td>
<td align="center">IMG</td>
<td align="center">Tooth</td>
<td align="center">Ursidae</td>
<td align="center">0.8</td>
<td align="center">0%</td>
<td align="center">&#x2014;</td>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
</tr>
<tr>
<td align="center">BRA-6115</td>
<td align="center">SG-CA-125</td>
<td align="center">Museum Collection</td>
<td align="center">IMG</td>
<td align="center">Tooth</td>
<td align="center">Ursidae</td>
<td align="center">0.5</td>
<td align="center">0%</td>
<td align="center">&#x2014;</td>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
</tr>
<tr>
<td align="center">BRA-6116</td>
<td align="center">SG-CA-10</td>
<td align="center">Museum Collection</td>
<td align="center">IMG</td>
<td align="center">Tooth</td>
<td align="center">Ursidae</td>
<td align="center">0.5</td>
<td align="center">0%</td>
<td align="center">&#x2014;</td>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
</tr>
<tr>
<td align="center">BRA-6117</td>
<td align="center">SG-CA-62</td>
<td align="center">Museum Collection</td>
<td align="center">IMG</td>
<td align="center">Bone</td>
<td align="center">Indeterminate</td>
<td align="center">0</td>
<td align="center">0%</td>
<td align="center">&#x2014;</td>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
</tr>
<tr>
<td align="center">BRA-6118</td>
<td align="center">SG-CA-137</td>
<td align="center">Museum Collection</td>
<td align="center">IMG</td>
<td align="center">Bone</td>
<td align="center">Caprini</td>
<td align="center">1.2</td>
<td align="center">0%</td>
<td align="center">&#x2014;</td>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
</tr>
<tr>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="center">Carbon (mg)</td>
<td align="center">Carbon (%)</td>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
</tr>
<tr>
<td align="center">BRA-6106</td>
<td align="center">SG21.11.53</td>
<td align="center">O300-PO400</td>
<td align="left">301-72</td>
<td align="center">Charcoal</td>
<td align="center">
<italic>Pinus sylvestris</italic> type</td>
<td align="center">33.3</td>
<td align="center">35%</td>
<td align="center">MAMS 56487</td>
<td align="center">&#x2212;22.61</td>
<td align="center">0.0009</td>
<td align="center">137.59</td>
<td align="center">&#x3e;49000</td>
<td align="center">&#x2014;</td>
<td align="center">&#x2014;</td>
<td align="center">&#x2014;</td>
</tr>
<tr>
<td align="center">BRA-6107</td>
<td align="center">SG21.03.05</td>
<td align="center">O300-PO400</td>
<td align="left">301-86</td>
<td align="center">Charcoal</td>
<td align="center">
<italic>Pinus sylvestris</italic> type</td>
<td align="center">50.8</td>
<td align="center">49%</td>
<td align="center">MAMS 56488</td>
<td align="center">&#x2212;23.43</td>
<td align="center">0.0012</td>
<td align="center">129.79</td>
<td align="center">&#x3e;49000</td>
<td align="center">&#x2014;</td>
<td align="center">&#x2014;</td>
<td align="center">&#x2014;</td>
</tr>
<tr>
<td align="center">BRA-6264</td>
<td align="center">SG-CA-200</td>
<td align="center">Museum Collection</td>
<td align="center">IMG</td>
<td align="center">Charcoal</td>
<td align="center">
<italic>Pinus sylvestris</italic> type</td>
<td align="center">4.6</td>
<td align="center">7%</td>
<td align="center">&#x2014;</td>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
<td align="left"/>
</tr>
<tr>
<td align="center">BRA-6265</td>
<td align="center">SG-CA-210</td>
<td align="center">Museum Collection</td>
<td align="center">IMG</td>
<td align="center">Charcoal</td>
<td align="center">
<italic>Pinus sylvestris</italic> type</td>
<td align="center">11.7</td>
<td align="center">21%</td>
<td align="center">MAMS 62623</td>
<td align="center">&#x2212;24.17</td>
<td align="center">0.0020</td>
<td align="center">87.05</td>
<td align="center">&#x3e;49000</td>
<td align="center">&#x2014;</td>
<td align="center">&#x2014;</td>
<td align="center">&#x2014;</td>
</tr>
<tr>
<td align="center">BRA-6266</td>
<td align="center">SG-CA-207</td>
<td align="center">Museum Collection</td>
<td align="center">IMG</td>
<td align="center">Charcoal</td>
<td align="center">
<italic>Pinus sylvestris</italic> type</td>
<td align="center">6.2</td>
<td align="center">16%</td>
<td align="center">MAMS 62624</td>
<td align="center">&#x2212;23.79</td>
<td align="center">0.0013</td>
<td align="center">172.17</td>
<td align="center">&#x3e;49000</td>
<td align="center">&#x2014;</td>
<td align="center">&#x2014;</td>
<td align="center">&#x2014;</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="s3-3">
<title>3.3 Paleoanthropology</title>
<p>A collection of 53 human remains was recovered by Mr. Aznar in 1978&#x2013;79, to which an upper third molar (SI-60) found <italic>in situ</italic> during the excavations (<xref ref-type="sec" rid="s10">Supplementary Table S2</xref>) must be added. The assemblage includes upper and lower dentition, a fragment of the mandible, vertebrae, the left and right upper limb (humeri and hand bones), and foot bones. Some of the specimens are preserved in complete and good condition, while others are fragmentary with old dry (diagenetic) and modern fractures (<xref ref-type="fig" rid="F2">Figure 2</xref>) (probably broken during the excavation). No evidence of pathological processes has been detected in the sample, while the preserved dentition does not display strong evidence of the use of the mouth in heavy-duty tasks.</p>
<fig id="F2" position="float">
<label>FIGURE 2</label>
<caption>
<p>Foremost Neanderthal remains found at the Simanya Gran gallery from Cova Simanya (Barcelona, Spain). Skeletal remains are distributed according to their attribution to specific individuals. <bold>(A)</bold> Left humerus; <bold>(B)</bold> shaft, caudal view; <bold>(C)</bold> bones of the foot; <bold>(D)</bold> bones of the right hand; <bold>(E)</bold> upper P<sup>3</sup> and M<sup>3</sup>; <bold>(F)</bold> fragment of the atlas and first phalanx of the hand; and <bold>(G)</bold> fragment of the ascending branch.</p>
</caption>
<graphic xlink:href="feart-11-1230707-g002.tif"/>
</fig>
<p>At least three individuals can be identified in the sample. The most complete one is defined by a complete left humerus SI-1 (<xref ref-type="fig" rid="F2">Figure 2A</xref>). Several other postcranial remains (right proximal humerus, hand 1 and hand 2, vertebra; <xref ref-type="fig" rid="F2">Figures 2B&#x2013;D</xref>) can be securely associated based on the size and taphonomic features. It corresponds to an adult, with a stature estimation for the humerus of 154 &#xb1; 4.89&#xa0;cm based on the <xref ref-type="bibr" rid="B37">Sj&#xf8;vold (1990)</xref> equation (max. length: 293&#xa0;mm), which is consistent with a Neanderthal female individual.</p>
<p>SI-7 is an upper left P3 at the Rc growth stage [according to the work of <xref ref-type="bibr" rid="B24">Moorrees et al. (1963)</xref>], with the root length completed with parallel ends (<xref ref-type="fig" rid="F2">Figure 2E</xref>). This provides an estimated age of approximately 11.5 years by modern human standards [according to the work of <xref ref-type="bibr" rid="B1">AlQahtani et al. (2010)</xref>]. SI-60 is a complete upper left M<sup>3</sup> crown (<xref ref-type="fig" rid="F2">Figure 2E</xref>) with a defined pulp roof [CRc, according to the work of <xref ref-type="bibr" rid="B24">Moorrees et al. (1963)</xref>], which would provide an age of 12.5&#x2013;13.5&#xa0;years [according to the work of <xref ref-type="bibr" rid="B1">AlQahtani et al. (2010)</xref>]. However, M3 development is thought to be accelerated in Neanderthals as compared to modern humans (<xref ref-type="bibr" rid="B18">Guatelli-Steinberg, 2009</xref>), which makes it probable that the M3 (SI-60) and P3 (SI-7) may come from a single individual of approximately 11.5&#xa0;years old.</p>
<p>Two other immature remains are also preserved in the sample: SI-29 is a hand first phalanx of ray V, whose size is like Juvenile 1 from El Sidr&#xf3;n (<xref ref-type="bibr" rid="B35">Rosas et al., 2017</xref>) (<xref ref-type="fig" rid="F2">Figure 2F</xref>). This provides an approximate age of 7.61&#x2013;7.78&#xa0;years for this individual. SI-37 is a fragment of an atlas from the right side, preserving upper and lower articular facets. The surface of the neurocentral junction can be appreciated, meaning that the anterior arch of the atlas was still unfused even though the size of the specimen is relatively large. This is fully compatible with the growth pattern established for Neanderthals by <xref ref-type="bibr" rid="B35">Rosas et al. (2017)</xref> and congruent with the age of 7.7 years established by the SI-37 phalanx.</p>
<p>The size of lower dentition I<sub>1</sub> (SI-3) and M<sub>3</sub> is relatively small, while P<sub>3</sub> (SI-5) falls at the upper limit and the upper I<sup>1</sup> (SI-45) in the middle of the Neanderthal distribution, while the mesiodistal crown diameter of the I<sup>2</sup> (SI-4 and SI-31) falls well above the Neanderthal mean [according to the work of <xref ref-type="bibr" rid="B28">Pinilla and Trinkaus (2017)</xref>] (<xref ref-type="sec" rid="s10">Supplementary Table S3</xref>, <xref ref-type="sec" rid="s10">Supplementary Figure S4</xref>). Historical and taphonomic conditions of the assemblage make it feasible to assign some of these teeth to individual 1. However, because of the noted size differences, the presence of other individuals cannot be ruled out. For the time being, no individual has been assigned to adult dentition. The upper and lower permanent teeth exhibit moderate occlusal wear [stage 5 of the work of <xref ref-type="bibr" rid="B38">Skinner (1997)</xref>], indicating a mature adult. Considering that Neanderthal anterior permanent teeth wear faster or commence wear earlier (<xref ref-type="bibr" rid="B38">Skinner, 1997</xref>), an age of 30&#x2013;40&#xa0;years may be considered.</p>
<p>In order to assess the location of the Simanya Gran specimen SI-1 in the morphospace (<xref ref-type="fig" rid="F3">Figure 3</xref>), we used 3D geometric morphometric techniques based on landmark configurations, according to <xref ref-type="bibr" rid="B34">Rosas et al.&#x2019;s (2015</xref>) procedures and comparative samples (see <xref ref-type="sec" rid="s10">Supplementary Material</xref>).</p>
<fig id="F3" position="float">
<label>FIGURE 3</label>
<caption>
<p>Scatter plot of the PCA (PC1&#x2013;PC2) shape space of the distal humerus performed on 18 landmarks and a large comparison sample from the work of <xref ref-type="bibr" rid="B34">Rosas et al. (2015)</xref>. Landmark 8 of the Simanya humerus has been estimated by the thin plate spline method. Ellipses for the prediction interval of a single new observation (0.95% probability) are shown only for the Neanderthal and modern human samples. Note the position of the Simanya&#x2019;s humerus clearly within the Neanderthal distribution.</p>
</caption>
<graphic xlink:href="feart-11-1230707-g003.tif"/>
</fig>
<p>Taxonomically, the human remains from Simanya Gran present a large series of Neanderthal apomorphic features distributed across different anatomical systems. Upper central incisors are large with a robust root, a strong dental tubercle, and characteristic flat wear (<xref ref-type="sec" rid="s10">Supplementary Figure S4</xref>). The upper lateral incisors present a conspicuous and characteristic shovel shape, where the marginal ridges are extremely thickened and invade the lingual fossa, circumscribing a narrow longitudinal groove. The mandible shows a large, moderately truncated gonial area, with a large medial pterygoid tubercle, and a lateral position of the condyle in relation to the mandibular notch (<xref ref-type="fig" rid="F2">Figure 2G</xref>). Humerus SI-1 (<xref ref-type="fig" rid="F2">Figures 2A</xref>, <xref ref-type="fig" rid="F3">3</xref>; <xref ref-type="sec" rid="s10">Supplementary Material</xref>) displays a massive and anteriorly projecting lesser tubercle. A deep olecranon fossa, which is elliptical in shape, wider than high, and a marked medial epicondyle are moderately prominent. The long lateral supracondylar crest and coracobrachial insertion are located more distally than the distal limit of the deltoid tuberosity, while the deltoid tuberosity displays two parallel and proximal ridges. The humeral torsion is moderate. The first metacarpal is short and displays a flange of bone along the distodorsoradial margin, marking the insertion of the opponens pollicis muscle. The first pollex phalanx is relatively short. The first phalanx of the second ray presents a robust and prominent radial tubercle, and the distal phalanges show an extremely wide distal end (<xref ref-type="fig" rid="F2">Figure 2D</xref>). The big toe metatarsal and distal phalanx are gracile, but the distal head of the metatarsal is proportionally bulky (<xref ref-type="fig" rid="F2">Figure 2C</xref>).</p>
<p>Ultimately, no positive results for aDNA were obtained, neither for shotgun sequencing nor for the mtDNA capture.</p>
</sec>
</sec>
<sec id="s4">
<title>4 Discussion and conclusion</title>
<p>Excavations at Simanya Gran revealed Pleistocene fossil-bearing deposits, primarily preserved in the layer O300-PO400 from the IMG and likely dismantled from the cave entrance. Its sub-superficial position aligns with Mr. Aznar&#x2019;s report, and similarities in the taxonomic structure and taphonomic features connect the &#x201c;Museum Collection&#x201d; to the documented contexts. The discovery of a Neanderthal upper M3 that is compatible with &#x201c;Museum Collection&#x201d; individual 2 supports this interpretation, which is further corroborated by Mr. Aznar identifying the excavation area.</p>
<p>The <sup>14</sup>C chronology provided by charcoals from our excavations are compatible with those from the charcoals included in the sediment samples associated to the human remains from the &#x201c;Museum Collection,&#x201d; potentially placing Simanya Neanderthals beyond 49,000&#xa0;years BP. A minimum age of <italic>circa</italic> 42,000&#xa0;cal BP has been obtained for the paleontological assemblage that is also present in the layer O300-PO400. The discrepancy between both ages is consistent with the open character of the layer that could have been deposited during a relatively long period including different time-segregated events.</p>
<p>The Simanya sample is attributed to Neanderthals due to distinct apomorphic traits (see <xref ref-type="fig" rid="F3">Figure 3</xref>). It includes at least three individuals&#x2014;an adult, a periadolescent (&#x223c;11.5&#xa0;years), and an immature (&#x223c;7.5&#xa0;years)&#x2014;with preserved dentition, mandible, vertebrae, upper limbs, and foot bones. Some specimens are complete and well preserved, while others are fragmentary with diagenetic and modern fractures. Taphonomic conditions allow assigning some teeth to individual 1, but based on size differences, a fourth individual is possible.</p>
<p>Simanya Neanderthals&#x2019; apomorphic traits align with other late Neanderthals, supporting, together with the chronological data from the excavation, its location in a final evolutionary phase of the evolution of western Neanderthals. However, different genetic backgrounds within Iberian Neanderthal populations cannot be ruled out. The substitution of Neanderthal populations in Western Europe (<xref ref-type="bibr" rid="B12">Dal&#xe9;n et al., 2012</xref>) and a population turnover with mtDNA diversity loss (&#x223c;100&#xa0;ka ago) have been identified (<xref ref-type="bibr" rid="B43">Vernot et al., 2021</xref>). The presence of primitive features in other samples, such as the 49,000 old from El Sidr&#xf3;n (<xref ref-type="bibr" rid="B33">Rosas et al., 2020</xref>), allows for hypothesizing the potential persistence of relict groups. Comparative studies with other Iberian samples may clarify late Iberian Neanderthals&#x2019; phylogeography.</p>
<p>The analysis of the Cova Simanya assemblage has allowed the identification of a new Neanderthal sample that will provide essential data on the morphological traits and evolutionary context of late Neanderthals in the Iberian Mediterranean region. Further comparative studies, incorporating morphometric and genetic analyses, with other Iberian samples will elucidate their phylogeography and population dynamics to shed light on a complex evolutionary history comprising different genetic backgrounds and relict genetic groups of late Iberian Neanderthals.</p>
</sec>
</body>
<back>
<sec sec-type="data-availability" id="s5">
<title>Data availability statement</title>
<p>The original contributions presented in the study are included in the article/<xref ref-type="sec" rid="s10">Supplementary Material</xref>; further inquiries can be directed to the corresponding authors.</p>
</sec>
<sec id="s6">
<title>Author contributions</title>
<p>Conceptualization: JM and AR. Investigation: JM, AC, MS, AR-H, RH, EM-R, DL, JR, DM-P, AG-T, EA, AG-B, EL, TM-B, ST, CL-F, and AR. Writing&#x2014;original draft: JM and AR. Writing&#x2014;review and editing: JM, AC, MS, AR-H, RH, EM-R, DL, JR, DM-P, AG-T, EA, AG-B, EL, TM-B, ST, CL-F, JF, and AR. Project administration: JM, JF, and AR. Funding acquisition: JM, JF, AR, MS, ST, and CL-F. All authors contributed to the article and approved the submitted version.</p>
</sec>
<sec id="s7">
<title>Funding</title>
<p>This research was funded by the Spanish Ministry of Science and Innovation through the projects PID2021-122356NB-I00 (MNCN-CSIC, AR), PID2020-113960GB-I00 (UB, JF), PID2021-124590NB-I00 (MCNB, CL-F), PID2019-103987GB-C31 (IPHES-CERCA), and PID2021-126004NB-100 (IBE-UPF-CSIC, TM-B); the AGAUR through the research groups 2021 SGR 01237 (IPHES-CERCA, EA), 2021 SGR 00337 (UB, JF), and 2021 SGR 00177 (IBE-UPF-CSIC, TM-B); the Culture Department of the Generalitat de Catalunya through the project ARQ001SOL-172-2022; and the Diputaci&#xf3; de Barcelona and the Fundaci&#xf3;n Palarq. MS was funded by the UAM Tom&#xe1;s y Valiente Program, JR by the European Union-Next Generation EU, Ministry of Universities and UA (MARSALAS21-22), and DL by the Xunta de Galicia Grant ED481B-2022-048. ST and TM-B received funding from the European Research Council under the European Union&#x2019;s Horizon 2020 Research and Innovation Program (grant agreement no. 803147 RESOLUTION, <ext-link ext-link-type="uri" xlink:href="https://site.unibo.it/resolution-erc/en">https://site.unibo.it/resolution-erc/en</ext-link> (ST), and no. 864203 (TM-B)]. IPHES-CERCA received financial support from the Spanish Ministry of Science and Innovation through the &#x201c;Mar&#xed;a de Maeztu&#x201d; program for Units of Excellence (CEX 2019-000945-M).</p>
</sec>
<ack>
<p>First and foremost, the authors would like to express their deepest gratitude to the Museu d&#x0027;Arqueologia de Catalunya for facilitating their access to the essential Aznar collection. The authors would also like to express their gratitude to the Parc Natural de Sant Lloren&#x00E7; del Munt i l&#x0027;Obac for its supportive role in facilitating the recommencement of excavations at Cova Simanya and for its commitment to the preservation of the site. They also extend their acknowledgement to Mr. Angel Mi&#x00F1;o and Mr. Josep Canals, whose assistance has been essential to their operations. Furthermore, they would like to acknowledge the consistent support provided by Mr. Joan Lleonart and Mr. Josep F&#x00ED;gols throughout the fieldwork. Their contributions have been beneficial to the progression and outcomes of the research.</p>
</ack>
<sec sec-type="COI-statement" id="s8">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec sec-type="disclaimer" id="s9">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors, and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec id="s10">
<title>Supplementary material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/feart.2023.1230707/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/feart.2023.1230707/full&#x23;supplementary-material</ext-link>
</p>
<supplementary-material xlink:href="DataSheet1.docx" id="SM1" mimetype="application/docx" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</sec>
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