Kent Academic Repository Full text document (pdf) Citation for published version Hublin, Jean-Jacques and Ben-Ncer, Abdelouahed and Bailey, Shara E. and Freidline, Sarah E. and Neubauer, Simon and Skinner, Matthew M. and Bergmann, Inga and Le Cabec, Adeline and Benazzi, Stefano and Harvati, Katerina and Gunz, Philipp (2017) New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens. Nature, 546 (7657). pp. 289-292. ISSN DOI https://doi.org/10.1038/nature22336 Link to record in KAR http://kar.kent.ac.uk/62267/ Document Version Author's Accepted Manuscript Copyright & reuse Content in the Kent Academic Repository is made available for research purposes. Unless otherwise stated all content is protected by copyright and in the absence of an open licence (eg Creative Commons), permissions for further reuse of content should be sought from the publisher, author or other copyright holder. 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Enquiries For any further enquiries regarding the licence status of this document, please contact: [email protected] If you believe this document infringes copyright then please contact the KAR admin team with the take-down information provided at http://kar.kent.ac.uk/contact.html 1 Title: Homo 2 sapiensNew fossils from Jebel )rhoud Morocco and the PanAfrican origin of 3 4 5 JeanJacques (ublin 6 Abdelouahed BenNcer 7 Shara E Bailey 8 Sarah E Freidline 9 Simon Neubauer 10 Matthew M Skinner 11 )nga Bergmann 12 Adeline Le Cabec 13 Stefano Benazzi 14 Katerina (arvati 15 Philipp Gunz 16 17 Department of (uman Evolution Max Planck )nstitute for Evolutionary Anthropology 18 Deutscher Platz Leipzig Germany 19 )nstitut National des Sciences de lArchéologie et du Patrimoine Rabat Morocco 20 Department of Anthropology Center for the Study of (uman Origins New York 21 University New York NY USA 22 School of Anthropology and Conservation University of Kent Canterbury CT NR 23 United Kingdom Department of Cultural (eritage University of Bologna Ravenna )taly 24 25 Paleoanthropology Senckenberg Center for (uman Evolution and Paleoenvironment 26 and DFG Center for Advanced Studies Words Bones Genes Tools Eberhard Karls Universität Tu bingen Germany 27 Fossil evidence points to an African origin of Homo sapiens from a group called 28 either H. heidelbergensis or H. rhodesiensis. However, the exact place and time of 29 our species’ emergence remain obscure because the fossil record is scarce and the 30 chronological age of many key specimens remains uncertain. In particular, it is 31 unclear whether the present day “modern” morphology emerged rapidly ca. 200 32 thousand years ago (ka) among earlier representatives of H. sapiens 1 or evolved 33 gradually over the last 400 ka2. Here, we report on new human fossils from Jebel 34 Irhoud (Morocco), and interpret the affinities of the hominins from this site with 35 other archaic and recent human groups. We identified a mosaic of features 36 including a facial, mandibular and dental morphology that aligns the Jebel Irhoud 37 material with early (EMH) or recent anatomically modern humans (RMH) and a 38 more primitive neurocranial and endocranial morphology. In combination with 39 the new date of 300-350 ka3, this evidence makes Jebel Irhoud the oldest and 40 richest African Middle Stone Age hominin site documenting early stages of the H. 41 sapiens clade in which key features of modern morphology were established. 42 Furthermore, it shows that the evolutionary processes behind the emergence of 43 our species were not restricted to sub-Saharan Africa. 44 45 46 )n mining operations in the Jebel )rhoud massif km southeast of Safi Morocco 47 exposed a Palaeolithic site in the Pleistocene filling of a karstic network An almost 48 complete skull )rhoud was accidentally unearthed in prompting excavations 49 that yielded an adult braincase )rhoud an immature mandible )rhoud an 50 immature humeral shaft an immature ilium and a fragment of mandible associated with abundant faunal remains and Levallois stone tool technology Although these 51 52 human remains were all reported to come from the bottom of the archaeological 53 deposits only the precise location of the humeral shaft was recorded 54 55 The interpretation of the )rhoud hominins has long been complicated by persistent 56 uncertainties surrounding their geological age They were initially assigned to a time 57 period ca ka ago and considered to be an African form of Neanderthal (owever 58 these affinities have been challenged and the faunal and microfaunal evidence 59 supported a middle Pleistocene MP age for the site An attempt to date one of the 60 hominins directly by U seriesESR suggested an age of ka Consistent with some genetic evidence fossils from Ethiopia Omo Kibish considered to be as old as 61 ~ ~ ka and (erto dated to ka are commonly regarded as the first EM( 62 63 )ntriguingly Omo and the (erto specimens appear to be more derived than the 64 supposedly contemporaneous or even younger )rhoud hominins )t has therefore been 65 H.suggested sapiens that the archaic features of the )rhoud fossils may indicate that North African 66 interbred with Neanderthals or that the )rhoud hominins represented a 67 North African late surviving archaic population 68 in situ 69 New excavations at )rhoud have recovered archaeological material and 70 established a precise chronology for the deposits which are much older than previously 71 thought The excavation yielded a new series of hominin remains including an adult 72 skull )rhoud comprising of a distorted braincase and fragments of the face Fig a 73 nearly complete adult mandible )rhoud Fig one maxilla several postcranial 74 elements and abundant dental material Extended Data Table These remains 75 primarily come from a single bone bed in the lower part of the archaeological deposits This concentration stratigraphical observations made by previous excavators and the 76 77 anatomical similarity with earlier discoveries strongly suggest that most if not all of the 78 hominin remains from the site were accumulated in a rather constrained window of 79 time corresponding to the formation of layer This layer contains the remains of at 80 least five individuals three adults one adolescent and one immature ca years old 81 )t now has a thermoluminescence TL weighted average age between and ka 82 with a probability compatible with a series of newly established U seriesESR 83 dates This timeframe places the )rhoud evidence in an entirely new perspective 84 Facial and mandibular morphology 85 86 When compared to the large robust and prognathic faces of the Neanderthals or older 87 MP forms the facial morphology of EM( and RM( is very distinctive The face is 88 relatively short and retracted under the braincase Facial structures are coronally 89 oriented and the infraorbital area is of inflexiontype displaying curvatures along the 90 horizontal sagittal and coronal profiles This pattern which may include some primitive 91 retentions strongly influences the morphology of the maxilla and zygomatic bone Our 92 morphometric analysis Fig and Methods clearly distinguishes MP archaic humans 93 and Neanderthals from RM( )n contrast all the possible reconstructions of the new 94 facial remains of )rhoud fall well within RM( variation as does )rhoud 95 96 Another facial characteristic observed in RM( is the weakness of their brow ridges 97 Some EM( from Africa and the Levant still display protruding supraorbital structures 98 but they tend to be dissociated into a medial superciliary arch and a lateral supraorbital 99 arch Among the )rhoud hominins these structures are rather variable which may be 100 related to sexual dimorphism )rhoud displays protruding supraorbital structures and the arches are poorly separated (owever in frontal view the supraorbital buttress 101 102 tends to form an inverted V above each orbit On )rhoud the torus is less projecting 103 and a modern pattern is already well expressed with a clear sulcus separating the two 104 arches On )rhoud the preserved parts do not display projecting supraorbital 105 structures Fig 106 107 The new )rhoud mandible is very large overall Fig Extended Data Table As in 108 some EM( from the Levant or North Africa it has retained a vertical symphysis with a 109 mental angleH. sapiens of Extended Data Fig The mandibular body displays a pattern 110 typical of its height strongly decreases from the front to the back This feature 111 is also present on the immature individual )rhoud Another modern aspect of )rhoud 112 is the rather narrow section of the mandibular body expressed by the breadthheight 113 index at the level of the mental foramen Extended Data Fig The )rhoud mandibles 114 also display some derived conditions in the mental area Extended Data Fig The 115 symphyseal section of )rhoud has a tearshaped outline quite distinctive of our 116 species Although the )rhoud mandibles lack a marked mandibular incurvation the 117 juvenile )rhoud displays a central keel between two depressions expanding inferiorlyH. 118 sapiensinto a thickened triangular eminence This inverted Tshape typical of recent 119 is incipient on the adult )ts inferior border is somewhat distended and 120 includes separated tubercles Notably this modern pattern is still inconsistently present 121 on Levantine EM( )n some aspects )rhoud is evocative of the Tabun mandible 122 but is much more robust 123 Dental morphology 124 125 The )rhoud teeth are generally very large Extended Data Tables and (owever their dental morphology is reminiscent of EM( in several respects The anterior teeth do 126 sapiens 127 not display the expansion observed in non MP hominins and Neanderthals and 128 the postcanine teeth are reduced compared to older hominins The M of the )rhoud 129 maxilla is already smaller than in some EM( The crown morphology Extended Data H.