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However, the scaphoid's complex morphology is challenging to comparatively analyze via traditional 2D linear measurements. This study quantifies scaphoid morphology utilizing 3D geometric morphometrics (3D GM) in African apes, humans and extinct hominins to provide a more holistic functional understanding of scaphoid morphology.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Materials and Methods<\/jats:title>\n                    <jats:p>\n                      We use 3D GM, including anatomical and sliding semi\u2010landmarks, to quantify scaphoid morphology in a comparative sample of African apes (\n                      <jats:styled-content style=\"fixed-case\">\n                        <jats:italic>Gorilla<\/jats:italic>\n                      <\/jats:styled-content>\n                      ,\n                      <jats:styled-content style=\"fixed-case\">\n                        <jats:italic>Gorilla beringei<\/jats:italic>\n                      <\/jats:styled-content>\n                      ,\n                      <jats:styled-content style=\"fixed-case\">\n                        <jats:italic>Pan paniscus<\/jats:italic>\n                      <\/jats:styled-content>\n                      and\n                      <jats:styled-content style=\"fixed-case\">\n                        <jats:italic>Pan troglodytes<\/jats:italic>\n                      <\/jats:styled-content>\n                      ) (\n                      <jats:italic>N<\/jats:italic>\n                      \u2009=\u200954), extant and fossil\n                      <jats:styled-content style=\"fixed-case\">\n                        <jats:italic>Homo sapiens<\/jats:italic>\n                      <\/jats:styled-content>\n                      (\n                      <jats:italic>N<\/jats:italic>\n                      \u2009=\u200920) and nine fossil hominin scaphoids from\n                      <jats:italic>Australopithecus<\/jats:italic>\n                      sp.,\n                      <jats:italic>Australopithecus sediba<\/jats:italic>\n                      ,\n                      <jats:italic>Homo naledi<\/jats:italic>\n                      and Neandertals.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      Principal component analysis indicates that extant species can be distinguished by differences in scaphoid shape that are consistent with variation in hand use during locomotion and manipulation. The australopith scaphoids plot between the African ape and modern human distributions, whereas\n                      <jats:italic>H. naledi<\/jats:italic>\n                      falls between\n                      <jats:italic>Gorilla<\/jats:italic>\n                      and human distributions.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Discussion<\/jats:title>\n                    <jats:p>\n                      Results confirm previous studies describing differences between extant African apes and modern human scaphoids that were interpreted as advantageous for knuckle\u2010walking and forceful manipulation, respectively. However, we highlight greater variation between\n                      <jats:italic>Pan<\/jats:italic>\n                      and\n                      <jats:italic>Gorilla<\/jats:italic>\n                      than previously recognized. The fossil hominin scaphoids present differing mosaics of joint orientation and shape, creating a distinct overall morphology in each hominin species. This may reflect differing functional pressures acting upon hominin wrists resulting from disparate combinations of locomotor and manipulative behaviors.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1002\/ajpa.70157","type":"journal-article","created":{"date-parts":[[2025,12,5]],"date-time":"2025-12-05T07:41:11Z","timestamp":1764920471000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Functional Morphology of the Scaphoid in Extant African Apes, Humans and Fossil Hominins"],"prefix":"10.1002","volume":"188","author":[{"ORCID":"https:\/\/orcid.org\/0009-0008-1598-6518","authenticated-orcid":false,"given":"Nadine G.","family":"Steer","sequence":"first","affiliation":[{"name":"School of Anthropology &amp; Conservation University of Kent  Canterbury UK"},{"name":"Department of Pathology and Anatomical Sciences University of Missouri  Columbia USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ameline","family":"Bardo","sequence":"additional","affiliation":[{"name":"D\u00e9partement Homme et Environnement Mus\u00e9e de l'Homme, UMR 7194 \u2010 HNHP, CNRS\u2010MNHN  Paris France"},{"name":"Department of Human Origins Max Planck Institute for Evolutionary Anthropology  Leipzig Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thomas W.","family":"Davies","sequence":"additional","affiliation":[{"name":"Department of Evolutionary Biology University of Vienna  Vienna Austria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5829-9952","authenticated-orcid":false,"given":"Antonio","family":"Rosas","sequence":"additional","affiliation":[{"name":"Paleoanthropology Group Museo Nacional de Ciencias Naturales\u2010Consejo Superior de Investigaciones Cient\u00edficas (CSIC)  Madrid Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Matthew M.","family":"Skinner","sequence":"additional","affiliation":[{"name":"Department of Human Origins Max Planck Institute for Evolutionary Anthropology  Leipzig Germany"},{"name":"Centre for the Exploration of the Deep Human Journey University of the Witwatersrand  Johannesburg South Africa"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5087-0897","authenticated-orcid":false,"given":"Tracy L.","family":"Kivell","sequence":"additional","affiliation":[{"name":"Department of Human Origins Max Planck Institute for Evolutionary Anthropology  Leipzig Germany"},{"name":"Centre for the Exploration of the Deep Human Journey University of the Witwatersrand  Johannesburg South Africa"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2025,12,4]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.jhevol.2023.103471"},{"key":"e_1_2_9_3_1","unstructured":"Adams D. 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