{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,30]],"date-time":"2026-04-30T00:37:49Z","timestamp":1777509469186,"version":"3.51.4"},"reference-count":77,"publisher":"Wiley","issue":"4","license":[{"start":{"date-parts":[[2017,4,21]],"date-time":"2017-04-21T00:00:00Z","timestamp":1492732800000},"content-version":"am","delay-in-days":365,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#am"},{"start":{"date-parts":[[2016,4,21]],"date-time":"2016-04-21T00:00:00Z","timestamp":1461196800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"}],"funder":[{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["DGE1255832"],"award-info":[{"award-number":["DGE1255832"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["American J Phys Anthropol"],"published-print":{"date-parts":[[2016,8]]},"abstract":"<jats:title>ABSTRACT<\/jats:title><jats:sec><jats:title>Objectives<\/jats:title><jats:p>Both interspecific and intraspecific variation in vertebral counts reflect the action of patterning control mechanisms such as <jats:italic>Hox<\/jats:italic>. The preserved A.L. 288\u20101 (\u201cLucy\u201d) sacrum contains five fused elements. However, the transverse processes of the most caudal element do not contact those of the segment immediately craniad to it, leaving incomplete sacral foramina on both sides. This conforms to the traditional definition of four\u2010segmented sacra, which are very rare in humans and African apes. It was recently suggested that fossilization damage precludes interpretation of this specimen and that additional sacral\u2010like features of its last segment (e.g., the extent of the sacral hiatus) suggest a general <jats:italic>Australopithecus<\/jats:italic> pattern of five sacral vertebrae.<\/jats:p><\/jats:sec><jats:sec><jats:title>Methods<\/jats:title><jats:p>We provide updated descriptions of the original Lucy sacrum. We evaluate sacral\/coccygeal variation in a large sample of extant hominoids and place it within the context of developmental variation in the mammalian vertebral column.<\/jats:p><\/jats:sec><jats:sec><jats:title>Results<\/jats:title><jats:p>We report that fossilization damage did not shorten the transverse processes of the fifth segment of Lucy's sacrum. In addition, we find that the extent of the sacral hiatus is too variable in apes and hominids<jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" xlink:href=\"#ajpa22997-note-0003\"\/> to provide meaningful information on segment identity. Most importantly, a combination of sacral and coccygeal features is to be expected in vertebrae at regional boundaries.<\/jats:p><\/jats:sec><jats:sec><jats:title>Discussion<\/jats:title><jats:p>The sacral\/caudal boundary appears to be displaced cranially in early hominids relative to extant African apes and humans, a condition consistent with the likely ancestral condition for Miocene hominoids. While not definitive in itself, a four\u2010segmented sacrum accords well with the \u201clong\u2010back\u201d model for the <jats:italic>Pan<\/jats:italic>\/<jats:italic>Homo<\/jats:italic> last common ancestor. Am J Phys Anthropol 160:729\u2013739, 2016. \u00a9 2016 Wiley Periodicals, Inc.<\/jats:p><\/jats:sec>","DOI":"10.1002\/ajpa.22997","type":"journal-article","created":{"date-parts":[[2016,4,21]],"date-time":"2016-04-21T17:04:28Z","timestamp":1461258268000},"page":"729-739","source":"Crossref","is-referenced-by-count":11,"title":["Developmental identity versus typology: Lucy has only four sacral segments"],"prefix":"10.1002","volume":"160","author":[{"given":"Allison L.","family":"Machnicki","sequence":"first","affiliation":[{"name":"Department of Anthropology Pennsylvania State University University Park PA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"C. Owen","family":"Lovejoy","sequence":"additional","affiliation":[{"name":"Department of Anthropology and School of Biomedical Sciences Kent State University Kent OH"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Philip L.","family":"Reno","sequence":"additional","affiliation":[{"name":"Department of Anthropology Pennsylvania State University University Park PA"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"311","published-online":{"date-parts":[[2016,4,21]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.cellbio.042308.113423"},{"key":"e_1_2_9_3_1","volume-title":"The Osteology of Infants and Children","author":"Baker BJ","year":"2005"},{"key":"e_1_2_9_4_1","doi-asserted-by":"publisher","DOI":"10.1006\/dbio.2000.9809"},{"key":"e_1_2_9_5_1","doi-asserted-by":"publisher","DOI":"10.1038\/316788a0"},{"key":"e_1_2_9_6_1","doi-asserted-by":"publisher","DOI":"10.1242\/dev.121.2.333"},{"key":"e_1_2_9_7_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2008.06.030"},{"key":"e_1_2_9_8_1","doi-asserted-by":"publisher","DOI":"10.1002\/bies.10088"},{"key":"e_1_2_9_9_1","doi-asserted-by":"publisher","DOI":"10.1038\/20944"},{"key":"e_1_2_9_10_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1107239"},{"key":"e_1_2_9_11_1","doi-asserted-by":"publisher","DOI":"10.1242\/dev.120.8.2187"},{"key":"e_1_2_9_12_1","doi-asserted-by":"publisher","DOI":"10.1016\/0092-8674(93)90378-4"},{"key":"e_1_2_9_13_1","doi-asserted-by":"publisher","DOI":"10.1242\/dev.001065"},{"key":"e_1_2_9_14_1","doi-asserted-by":"publisher","DOI":"10.1093\/molehr\/3.2.115"},{"key":"e_1_2_9_15_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.92.1.310"},{"key":"e_1_2_9_16_1","doi-asserted-by":"publisher","DOI":"10.1242\/dev.122.2.449"},{"key":"e_1_2_9_17_1","first-page":"414","article-title":"On the completion of ossification of the human sacrum","volume":"30","author":"Fawcett E.","year":"1907","journal-title":"Anat Anz"},{"key":"e_1_2_9_18_1","doi-asserted-by":"publisher","DOI":"10.1242\/dev.122.2.461"},{"key":"e_1_2_9_19_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0016-6995(02)00022-0"},{"key":"e_1_2_9_20_1","doi-asserted-by":"publisher","DOI":"10.1006\/jhev.2002.0595"},{"key":"e_1_2_9_21_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.jhevol.2011.07.004"},{"key":"e_1_2_9_22_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.jhevol.2012.05.011"},{"key":"e_1_2_9_23_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0070-2153(09)88007-1"},{"key":"e_1_2_9_24_1","doi-asserted-by":"publisher","DOI":"10.1002\/j.1460-2075.1991.tb07764.x"},{"key":"e_1_2_9_25_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajpa.1330570403"},{"key":"e_1_2_9_26_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0901280106"},{"key":"e_1_2_9_27_1","doi-asserted-by":"publisher","DOI":"10.1002\/bies.950140408"},{"key":"e_1_2_9_28_1","doi-asserted-by":"publisher","DOI":"10.1002\/(SICI)1520-6505(1998)6:3<87::AID-EVAN3>3.0.CO;2-T"},{"key":"e_1_2_9_29_1","doi-asserted-by":"publisher","DOI":"10.1002\/(SICI)1096-8644(1999)110:29 <189::AID-AJPA7>3.0.CO;2-3"},{"key":"e_1_2_9_30_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.279.5359.2115"},{"key":"e_1_2_9_31_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.gaitpost.2004.06.010"},{"key":"e_1_2_9_32_1","first-page":"95","article-title":"The natural history of human gait and posture. Part 1: Spine and pelvis","volume":"21","author":"Lovejoy CO.","year":"2005","journal-title":"Gait Posture"},{"key":"e_1_2_9_33_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.gaitpost.2006.05.001"},{"key":"e_1_2_9_34_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1175832"},{"key":"e_1_2_9_35_1","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.2010.0112"},{"key":"e_1_2_9_36_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1175827"},{"key":"e_1_2_9_37_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1175833"},{"key":"e_1_2_9_38_1","doi-asserted-by":"publisher","DOI":"10.7717\/peerj.1521"},{"key":"e_1_2_9_39_1","doi-asserted-by":"publisher","DOI":"10.1006\/jhev.2002.0554"},{"key":"e_1_2_9_40_1","doi-asserted-by":"publisher","DOI":"10.1387\/ijdb.072276mm"},{"key":"e_1_2_9_41_1","doi-asserted-by":"publisher","DOI":"10.1002\/jez.b.21316"},{"key":"e_1_2_9_42_1","doi-asserted-by":"publisher","DOI":"10.1242\/dev.007567"},{"key":"e_1_2_9_43_1","doi-asserted-by":"publisher","DOI":"10.1006\/dbio.2000.9683"},{"key":"e_1_2_9_44_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1420275111"},{"key":"e_1_2_9_45_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0912622107"},{"key":"e_1_2_9_46_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7580.2005.00473.x"},{"key":"e_1_2_9_47_1","doi-asserted-by":"publisher","DOI":"10.1002\/jez.b.22"},{"key":"e_1_2_9_48_1","doi-asserted-by":"publisher","DOI":"10.1038\/348237a0"},{"key":"e_1_2_9_49_1","doi-asserted-by":"publisher","DOI":"10.1002\/evan.21417"},{"key":"e_1_2_9_50_1","doi-asserted-by":"publisher","DOI":"10.1002\/jez.b.22506"},{"key":"e_1_2_9_51_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.92.18.8185"},{"key":"e_1_2_9_52_1","volume-title":"Cunningham's Text\u2010Book of Anatomy","author":"Robinson A.","year":"1913"},{"key":"e_1_2_9_53_1","first-page":"234","volume-title":"Triangulating the Evolution of the Vertebral Column in the Last Common Ancestor: Thoracolumbar Transverse Process Homology in the Hominoidea","author":"Rosenman BA.","year":"2008"},{"key":"e_1_2_9_54_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajpa.22642"},{"key":"e_1_2_9_55_1","doi-asserted-by":"publisher","DOI":"10.1038\/374460a0"},{"key":"e_1_2_9_56_1","first-page":"601","article-title":"The numbers of vertebrae in primates","volume":"89","author":"Schultz AH","year":"1945","journal-title":"Proc Am Phil Soc"},{"key":"e_1_2_9_57_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0604706103"},{"key":"e_1_2_9_58_1","doi-asserted-by":"publisher","DOI":"10.1101\/gad.7.12a.2318"},{"key":"e_1_2_9_59_1","doi-asserted-by":"publisher","DOI":"10.1101\/gad.205701"},{"key":"e_1_2_9_60_1","volume-title":"Gray's Anatomy","author":"Standring S.","year":"2005"},{"key":"e_1_2_9_61_1","doi-asserted-by":"publisher","DOI":"10.1093\/molbev\/msn218"},{"key":"e_1_2_9_62_1","doi-asserted-by":"publisher","DOI":"10.1006\/dbio.2000.9651"},{"key":"e_1_2_9_63_1","doi-asserted-by":"publisher","DOI":"10.1016\/0925-4773(95)00371-1"},{"key":"e_1_2_9_64_1","doi-asserted-by":"publisher","DOI":"10.1101\/gad.1547407"},{"key":"e_1_2_9_65_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrg2099"},{"key":"e_1_2_9_66_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-662-10382-1_10"},{"key":"e_1_2_9_67_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-642-39979-4_34"},{"key":"e_1_2_9_68_1","doi-asserted-by":"publisher","DOI":"10.1002\/dvdy.21286"},{"key":"e_1_2_9_69_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0070-2153(09)88009-5"},{"key":"e_1_2_9_70_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1085672"},{"key":"e_1_2_9_71_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1078294"},{"key":"e_1_2_9_72_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1403659111"},{"key":"e_1_2_9_73_1","unstructured":"WilliamsSA.2011a. Evolution of the Hominoid Vertebral Column: Universtiy of Illinois at Urbana\u2010Champaign.330p."},{"key":"e_1_2_9_74_1","doi-asserted-by":"crossref","first-page":"134","DOI":"10.1002\/jezb.21451","article-title":"Variation in anthropoid vertebral formulae: implications for homology and homoplasy in hominoid evolution","volume":"318","author":"Williams SA.","year":"2011","journal-title":"J Exp Zool (Mol Dev Evol)"},{"key":"e_1_2_9_75_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajpa.22049"},{"key":"e_1_2_9_76_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1232996"},{"key":"e_1_2_9_77_1","doi-asserted-by":"publisher","DOI":"10.1002\/evan.21437"},{"key":"e_1_2_9_78_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00586-012-2595-2"}],"container-title":["American Journal of Physical Anthropology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.wiley.com\/onlinelibrary\/tdm\/v1\/articles\/10.1002%2Fajpa.22997","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.wiley.com\/onlinelibrary\/tdm\/v1\/articles\/10.1002%2Fajpa.22997","content-type":"unspecified","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/ajpa.22997","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1002\/ajpa.22997","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/am-pdf\/10.1002\/ajpa.22997","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/ajpa.22997","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,9,13]],"date-time":"2023-09-13T06:49:16Z","timestamp":1694587756000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1002\/ajpa.22997"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2016,4,21]]},"references-count":77,"journal-issue":{"issue":"4","published-print":{"date-parts":[[2016,8]]}},"alternative-id":["10.1002\/ajpa.22997"],"URL":"https:\/\/doi.org\/10.1002\/ajpa.22997","archive":["Portico"],"relation":{},"ISSN":["0002-9483","1096-8644"],"issn-type":[{"value":"0002-9483","type":"print"},{"value":"1096-8644","type":"electronic"}],"subject":[],"published":{"date-parts":[[2016,4,21]]}}}