{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T22:28:01Z","timestamp":1792794481706,"version":"4.5.0"},"reference-count":92,"publisher":"Public Library of Science (PLoS)","issue":"1","license":[{"start":{"date-parts":[[2026,1,29]],"date-time":"2026-01-29T00:00:00Z","timestamp":1769644800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100004871","name":"Research Foundation for the State University of New York","doi-asserted-by":"publisher","award":["N\/A"],"award-info":[{"award-number":["N\/A"]}],"id":[{"id":"10.13039\/100004871","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000038","name":"Natural Sciences and Engineering Research Council of Canada","doi-asserted-by":"publisher","award":["RGPIN-2020-04159"],"award-info":[{"award-number":["RGPIN-2020-04159"]}],"id":[{"id":"10.13039\/501100000038","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.plosone.org"],"crossmark-restriction":false},"short-container-title":["PLoS One"],"abstract":"<jats:p>Humans are unique among primates in possessing a chin, yet it is currently unclear whether the form of the symphyseal region of the mandible where the chin is located is the product of direct selection or a by-product of evolutionary pressures on other craniomandibular features. Here, we conduct an evolutionary analysis of hominoid craniomandibular traits to test three hypotheses: symphyseal mandibular traits evolved (1) neutrally due to genetic drift, (2) under direct selection, and (3) as a by-product (or \u201cspandrel\u201d) of selection on other craniomandibular traits. Evolutionary rates of morphological change, via Lande\u2019s generalized genetic distance, were estimated along each branch of a fully-resolved hominoid phylogeny to reveal patterns of neutral, stabilizing and directional selection. Directional selection was detected along the branch between humans and the last common ancestor of chimpanzees and humans, against a backdrop of pervasive stabilizing selection and neutral evolution in hominoids. Significant directional selection was found on cranial traits reflecting increased basicranial flexion, neurocranial expansion, and reduction in lower facial prognathism, and on mandibular traits that generate a more parabolic-shaped, gracile mandible with a smaller ramus and shallower corpus. In contrast, of the nine mandibular \u201cchin\u201d traits, only three were under significant direct selection, while the other six were either under no selection or indirect selection. Thus, the results are consistent with the hypothesis that the symphyseal morphology that forms the human chin evolved largely as a by-product (i.e., spandrel) of direct selection for reduced anterior dental size and the craniofacial changes correlated with the evolution of bipedalism in hominins, rather than as a specific adaptation.<\/jats:p>","DOI":"10.1371\/journal.pone.0340278","type":"journal-article","created":{"date-parts":[[2026,1,29]],"date-time":"2026-01-29T18:29:27Z","timestamp":1769711367000},"page":"e0340278","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":1,"title":["Is the human chin a spandrel? Insights from an evolutionary analysis of ape craniomandibular form"],"prefix":"10.1371","volume":"21","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5263-3892","authenticated-orcid":true,"given":"Noreen","family":"von Cramon-Taubadel","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jill E.","family":"Scott","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8240-9915","authenticated-orcid":true,"given":"Chris A.","family":"Robinson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lauren","family":"Schroeder","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2026,1,29]]},"reference":[{"issue":"4","key":"pone.0340278.ref001","doi-asserted-by":"crossref","first-page":"247","DOI":"10.1006\/jhev.1993.1048","article-title":"The relationship of in vivo bone strain to mandibular corpus morphology in Macaca fascicularis","volume":"25","author":"DJ Daegling","year":"1993","journal-title":"Journal of Human Evolution"},{"issue":"1","key":"pone.0340278.ref002","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1002\/evan.21471","article-title":"The enduring puzzle of the human chin","volume":"25","author":"JD Pampush","year":"2016","journal-title":"Evol Anthropol"},{"issue":"706","key":"pone.0340278.ref003","first-page":"439","article-title":"The evolution of the chin","volume":"200","author":"L Robinson","year":"1914","journal-title":"The North American Review"},{"issue":"3","key":"pone.0340278.ref004","doi-asserted-by":"crossref","first-page":"367","DOI":"10.1006\/jhev.1999.0339","article-title":"The human chin revisited: what is it and who has it?","volume":"38","author":"JH Schwartz","year":"2000","journal-title":"J Hum Evol"},{"key":"pone.0340278.ref005","first-page":"1","article-title":"The mandibles of Sinanthropus pekinensis: a comparative study","volume":"7","author":"F Weidenreich","year":"1936","journal-title":"Paleontologia Sinica, Ser D"},{"issue":"2","key":"pone.0340278.ref006","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1002\/jez.b.23249","article-title":"What is a trait? 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Cramon-Taubadel","year":"2012","journal-title":"J Hum Evol"},{"issue":"1","key":"pone.0340278.ref037","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1002\/ajpa.20616","article-title":"The problem of assessing landmark error in geometric morphometrics: theory, methods, and modifications","volume":"134","author":"N von Cramon-Taubadel","year":"2007","journal-title":"Am J Phys Anthropol"},{"key":"pone.0340278.ref038","doi-asserted-by":"crossref","first-page":"94","DOI":"10.1016\/j.jhevol.2015.05.008","article-title":"Complex and changing patterns of natural selection explain the evolution of the human hip","volume":"85","author":"M Grabowski","year":"2015","journal-title":"J Hum Evol"},{"issue":"1","key":"pone.0340278.ref039","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1006\/mpev.1996.0056","article-title":"Mitochondrial DNA diversity in gorillas","volume":"6","author":"KJ Garner","year":"1996","journal-title":"Mol Phylogenet Evol"},{"issue":"12","key":"pone.0340278.ref040","doi-asserted-by":"crossref","first-page":"4766","DOI":"10.1073\/pnas.1015422108","article-title":"Evidence from Cameroon reveals differences in the genetic structure and histories of chimpanzee populations","volume":"108","author":"MK Gonder","year":"2011","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"3","key":"pone.0340278.ref041","doi-asserted-by":"crossref","first-page":"447","DOI":"10.1016\/j.ympev.2010.11.005","article-title":"Unresolved molecular phylogenies of gibbons and siamangs (Family: Hylobatidae) based on mitochondrial, Y-linked, and X-linked loci indicate a rapid Miocene radiation or sudden vicariance event","volume":"58","author":"H Israfil","year":"2011","journal-title":"Mol Phylogenet Evol"},{"issue":"19","key":"pone.0340278.ref042","doi-asserted-by":"crossref","first-page":"8900","DOI":"10.1073\/pnas.91.19.8900","article-title":"Gene trees and hominoid phylogeny","volume":"91","author":"M Ruvolo","year":"1994","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"6","key":"pone.0340278.ref043","doi-asserted-by":"crossref","first-page":"1699","DOI":"10.1111\/j.1558-5646.1997.tb05095.x","article-title":"LIKELIHOOD OF ANCESTOR STATES IN ADAPTIVE RADIATION","volume":"51","author":"D Schluter","year":"1997","journal-title":"Evolution"},{"issue":"1","key":"pone.0340278.ref044","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1111\/bij.12765","article-title":"A multiple variance Brownian motion framework for estimating variable rates and inferring ancestral states","volume":"118","author":"JB Smaers","year":"2016","journal-title":"Biol J Linn Soc"},{"issue":"5","key":"pone.0340278.ref045","doi-asserted-by":"crossref","first-page":"1341","DOI":"10.1111\/j.1558-5646.1997.tb01457.x","article-title":"Stabilizing selection and the comparative analysis of adaptation","volume":"51","author":"TF Hansen","year":"1997","journal-title":"Evolution"},{"issue":"5","key":"pone.0340278.ref046","doi-asserted-by":"crossref","first-page":"1201","DOI":"10.1111\/j.1420-9101.2008.01573.x","article-title":"Measuring and comparing evolvability and constraint in multivariate characters","volume":"21","author":"TF Hansen","year":"2008","journal-title":"J Evol Biol"},{"issue":"3","key":"pone.0340278.ref047","doi-asserted-by":"crossref","first-page":"304","DOI":"10.2307\/2992324","article-title":"Squared-Change Parsimony Reconstructions of Ancestral States for Continuous-Valued Characters on a Phylogenetic Tree","volume":"40","author":"WP Maddison","year":"1991","journal-title":"Systematic Zoology"},{"issue":"2","key":"pone.0340278.ref048","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1093\/bioinformatics\/btg412","article-title":"APE: Analyses of Phylogenetics and Evolution in R language","volume":"20","author":"E Paradis","year":"2004","journal-title":"Bioinformatics"},{"issue":"5","key":"pone.0340278.ref049","doi-asserted-by":"crossref","first-page":"958","DOI":"10.1111\/j.1558-5646.1988.tb02514.x","article-title":"A comparison of genetic and phenotypic correlations","volume":"42","author":"JM Cheverud","year":"1988","journal-title":"Evolution"},{"issue":"2","key":"pone.0340278.ref050","doi-asserted-by":"crossref","first-page":"186","DOI":"10.1017\/pab.2021.36","article-title":"Evolvability in the fossil record","volume":"48","author":"AC Love","year":"2021","journal-title":"Paleobiology"},{"issue":"3","key":"pone.0340278.ref051","doi-asserted-by":"crossref","first-page":"260","DOI":"10.1002\/ajpa.10038","article-title":"Discerning evolutionary processes in patterns of tamarin (genus Saguinus) craniofacial variation","volume":"117","author":"RR Ackermann","year":"2002","journal-title":"Am J Phys Anthropol"},{"issue":"4","key":"pone.0340278.ref052","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1016\/j.jhevol.2009.01.010","article-title":"Covariance structure in the skull of Catarrhini: a case of pattern stasis and magnitude evolution","volume":"56","author":"FB de Oliveira","year":"2009","journal-title":"J Hum Evol"},{"issue":"1","key":"pone.0340278.ref053","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1007\/s11692-008-9038-3","article-title":"The Evolution of Modularity in the Mammalian Skull I: Morphological Integration Patterns and Magnitudes","volume":"36","author":"A Porto","year":"2008","journal-title":"Evol Biol"},{"issue":"4","key":"pone.0340278.ref054","doi-asserted-by":"crossref","first-page":"293","DOI":"10.1007\/s11692-020-09514-w","article-title":"Examination of Sample Size Determination in Integration Studies Based on the Integration Coefficient of Variation (ICV)","volume":"47","author":"H Jung","year":"2020","journal-title":"Evol 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Sjovold","year":"1984"},{"issue":"6648","key":"pone.0340278.ref060","doi-asserted-by":"crossref","first-page":"913","DOI":"10.1126\/science.abn6919","article-title":"Phylogenomic analyses provide insights into primate evolution","volume":"380","author":"Y Shao","year":"2023","journal-title":"Science"},{"issue":"7459","key":"pone.0340278.ref061","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1038\/nature12228","article-title":"Great ape genetic diversity and population history","volume":"499","author":"J Prado-Martinez","year":"2013","journal-title":"Nature"},{"issue":"6","key":"pone.0340278.ref062","doi-asserted-by":"crossref","first-page":"1210","DOI":"10.1111\/j.1558-5646.1983.tb00236.x","article-title":"The measurement of selection on correlated characters","volume":"37","author":"R Lande","year":"1983","journal-title":"Evolution"},{"issue":"2","key":"pone.0340278.ref063","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1093\/icb\/23.2.347","article-title":"Morphology, Performance and Fitness","volume":"23","author":"SJ Arnold","year":"1983","journal-title":"Am Zool"},{"issue":"1","key":"pone.0340278.ref064","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1146\/annurev.an.11.100182.001055","article-title":"The Pattern of Human Evolution: Studies on Bipedalism, Mastication, and Encephalization","volume":"11","author":"HM McHenry","year":"1982","journal-title":"Annu Rev Anthropol"},{"issue":"6","key":"pone.0340278.ref065","doi-asserted-by":"crossref","first-page":"661","DOI":"10.1002\/jmor.20246","article-title":"Ontogenetic variation in the mandibular ramus of great apes and humans","volume":"275","author":"CE Terhune","year":"2014","journal-title":"J 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Soc"},{"issue":"1","key":"pone.0340278.ref084","doi-asserted-by":"crossref","first-page":"44","DOI":"10.1093\/icb\/36.1.44","article-title":"Developmental Integration and the Evolution of Pleiotropy","volume":"36","author":"JM Cheverud","year":"1996","journal-title":"Am Zool"},{"issue":"5","key":"pone.0340278.ref085","doi-asserted-by":"crossref","first-page":"359","DOI":"10.1016\/j.jhevol.2004.08.009","article-title":"Facial heights: evolutionary relevance of postnatal ontogeny for facial orientation and skull morphology in humans and chimpanzees","volume":"47","author":"M Bastir","year":"2004","journal-title":"J Hum Evol"},{"issue":"9","key":"pone.0340278.ref086","doi-asserted-by":"crossref","first-page":"814","DOI":"10.1016\/j.archoralbio.2006.03.009","article-title":"Correlated variation between the lateral basicranium and the face: a geometric morphometric study in different human groups","volume":"51","author":"M Bastir","year":"2006","journal-title":"Arch Oral Biol"},{"key":"pone.0340278.ref087","doi-asserted-by":"crossref","first-page":"103256","DOI":"10.1016\/j.jhevol.2022.103256","article-title":"Morphological modularity in the anthropoid axial skeleton","volume":"172","author":"H Jung","year":"2022","journal-title":"J Hum Evol"},{"issue":"1","key":"pone.0340278.ref088","doi-asserted-by":"crossref","first-page":"84","DOI":"10.1002\/ajpa.23606","article-title":"Neuromandibular integration in humans and chimpanzees: Implications for dental and mandibular reduction in Homo","volume":"167","author":"A Veneziano","year":"2018","journal-title":"Am J Phys Anthropol"},{"issue":"5","key":"pone.0340278.ref089","doi-asserted-by":"crossref","first-page":"637","DOI":"10.1111\/j.1469-7580.2006.00644.x","article-title":"Craniofacial levels and the morphological maturation of the human skull","volume":"209","author":"M Bastir","year":"2006","journal-title":"J 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