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Hublin, Jean-Jacques, Ben-Ncer, Abdelouahed, Bailey, Shara E., Freidline,
Sarah E., Neubauer, Simon, Skinner, Matthew M., Bergmann, Inga, Le Cabec,
Adeline, Benazzi, Stefano, Harvati, Katerina and and others (2017) New
fossils from Jebel Irhoud, Morocco and the pan-African origin of Homosapiens. Nature, 546 (7657). pp. 289-292. ISSN 0028-0836.

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1

Title: New fossils from Jebel Irhoud (Morocco) and the Pan-African origin of Homo

2

sapiens.

3
4Jean-Jacques Hublin1

5Abdelouahed Ben-Ncer2

6

Shara E. Bailey3

7

Sarah E. Freidline1

8

Simon Neubauer1

9Matthew M. Skinner4

10Inga Bergmann1

11

Adeline Le Cabec1

12

Stefano Benazzi5

13

Katerina Harvati6

14Philipp Gunz1

15
16

1 Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology,

17

Deutscher Platz 6, Leipzig, 04103, Germany

18

2 Institut National des Sciences de l'Archéologie et du Patrimoine, Rabat, Morocco19

3Department of Anthropology, Center for the Study of Human Origins, New York

20

University, New York, NY 10003, USA

21

4School of Anthropology and Conservation, University of Kent, Canterbury CT2 7NR,

22

United Kingdom

23

5Department of Cultural Heritage, University of Bologna, Ravenna 48121, Italy

24

6Paleoanthropology, Senckenberg Center for Human Evolution and Paleoenvironment,

25

and DFG Center for Advanced Studies: “Words, Bones, Genes, Tools”, Eberhard Karls

26

Universität, Tü bingen, Germany

27

Fossil evidence points to an African origin of Homo sapiens from a group called28

either H. heidelbergensis or H. rhodesiensis. However, the exact place and time of29

our species’ emergence remain obscure because the fossil record is scarce and the30chronological age of many key specimens remains uncertain. In particular, it is31

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 with35

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 a38

more primitive neurocranial and endocranial morphology. In combination with39

the new date of 300-350 ka3, this evidence makes Jebel Irhoud the oldest and40richest African Middle Stone Age hominin site documenting early stages of the H.

41sapiens clade in which key features of modern morphology were established.

42

Furthermore, it shows that the evolutionary processes behind the emergence of43

our species were not restricted to sub-Saharan Africa.

44
45

In 1960, mining operations in the Jebel Irhoud massif 55 km southeast of Safi, Morocco

46

exposed a Palaeolithic site in the Pleistocene filling of a karstic network. An almost

47

complete skull (Irhoud 1) was accidentally unearthed in 1961, prompting excavations

48

that yielded an adult braincase (Irhoud 2)4, an immature mandible (Irhoud 3)5, an

49

immature humeral shaft6, an immature ilium7 and a fragment of mandible8 associated50with abundant faunal remains and Levallois stone tool technology6. Although these51

human remains were all reported to come from the bottom of the archaeological52

deposits, only the precise location of the humeral shaft was recorded.

53
54

The interpretation of the Irhoud hominins has long been complicated by persistent55

uncertainties surrounding their geological age. They were initially assigned to a time56

period ca. 40 ka ago and considered to be an African form of Neanderthal9. However,57

these affinities have been challenged5,10,11 and the faunal8 and microfaunal12 evidence

58

supported a middle Pleistocene (MP) age for the site. An attempt to date one of the59

hominins directly by U series-ESR3 suggested an age of 160 ± 16 ka13. Consistent with60

some genetic evidence14, fossils from Ethiopia (Omo Kibish, considered to be as old as

61~195 ka15 and Herto, dated to ~16016 ka) are commonly regarded as the first EMH.

62

Intriguingly, Omo 1 and the Herto specimens appear to be more derived than the

63

supposedly contemporaneous or even younger Irhoud hominins. It has therefore been64

suggested that the archaic features of the Irhoud fossils may indicate that North African65

H. sapiens interbred with Neanderthals17, or that the Irhoud hominins represented a

66

North African, late surviving, archaic population18.

67
68

New excavations at Irhoud have recovered in situ archaeological material and69

established a precise chronology for the deposits, which are much older than previously

70

thought3. The excavation yielded a new series of hominin remains, including an adult71

skull (Irhoud 10) comprising of a distorted braincase and fragments of the face (Fig. 1), a

72

nearly complete adult mandible (Irhoud 11) (Fig. 2), one maxilla, several postcranial

73

elements, and abundant dental material (Extended Data Table 1). These remains

74

primarily come from a single bone bed in the lower part of the archaeological deposits.

75

This concentration, stratigraphical observations made by previous excavators, and the

76

anatomical similarity with earlier discoveries strongly suggest that most, if not all, of the77

hominin remains from the site were accumulated in a rather constrained window of

78

time corresponding to the formation of layer 7. This layer contains the remains of at

79

least five individuals (three adults, one adolescent and one immature, ca. 7.5 years old).

80

It now has a thermoluminescence (TL) weighted average age between 300 and 350 ka

81with a 95 % probability, compatible with a series of newly established U series-ESR

82

dates3. This timeframe places the Irhoud evidence in an entirely new perspective.

83
84

Facial and mandibular morphology

85

When compared to the large, robust and prognathic faces of the Neanderthals or older86

MP forms, the facial morphology of EMH and RMH is very distinctive. The face is87relatively short and retracted under the braincase. Facial structures are coronally88

oriented and the infraorbital area is of “inflexion-type”, displaying curvatures along the89

horizontal, sagittal and coronal profiles. This pattern, which may include some primitive90

retentions19, strongly influences the morphology of the maxilla and zygomatic bone. Our

91morphometric analysis (Fig. 3 and Methods) clearly distinguishes MP archaic humans

92

and Neanderthals from RMH. In contrast, all the possible reconstructions of the new93

facial remains of Irhoud 10 fall well within RMH variation, as does Irhoud 1.

94
95

Another facial characteristic observed in RMH is the weakness of their brow ridges.

96

Some EMH from Africa and the Levant still display protruding supraorbital structures,

97

but they tend to be dissociated into a medial superciliary arch and a lateral supraorbital

98

arch. Among the Irhoud hominins these structures are rather variable, which may be99

related to sexual dimorphism. Irhoud 1 displays protruding supraorbital structures and100

the arches are poorly separated. However in frontal view, the supraorbital buttress101

tends to form an inverted V above each orbit. On Irhoud 2, the torus is less projecting

102

and a modern pattern is already well expressed, with a clear sulcus separating the two103

arches. On Irhoud 10, the preserved parts do not display projecting supraorbital

104

structures (Fig. 1).

105
106

The new Irhoud 11 mandible is very large overall (Fig. 2, Extended Data Table 2). As in

107

some EMH from the Levant or North Africa, it has retained a vertical symphysis, with a108

mental angle of 88.8° (Extended Data Fig. 1). The mandibular body displays a pattern

109

typical of H. sapiens: its height strongly decreases from the front to the back. This feature110

is also present on the immature individual Irhoud 3. Another modern aspect of Irhoud111

11 is the rather narrow section of the mandibular body expressed by the breadth/height112

index at the level of the mental foramen (Extended Data Fig. 1). The Irhoud mandibles

113

also display some derived conditions in the mental area (Extended Data Fig. 1). The

114

symphyseal section of Irhoud 11 has a tear-shaped outline quite distinctive of our115

species. Although the Irhoud mandibles lack a marked mandibular incurvation, the116

juvenile Irhoud 3 displays a central keel between two depressions expanding inferiorly

117

into a thickened triangular eminence. This inverted T-shape, typical of recent H.

118

sapiens20, is incipient on the adult. Its inferior border is somewhat distended and119

includes separated tubercles. Notably, this modern pattern is still inconsistently present

120

on Levantine EMH20. In some aspects, Irhoud 11 is evocative of the Tabun 2 mandible,

121

but is much more robust.

122
123

Dental morphology

124

The Irhoud teeth are generally very large (Extended Data Tables 3 and 4). However,125

their dental morphology is reminiscent of EMH in several respects. The anterior teeth do

126

not display the expansion observed in non-sapiens MP hominins and Neanderthals21 and127

the postcanine teeth are reduced compared to older hominins. The M3 of the Irhoud 21

128

maxilla is already smaller than in some EMH. The crown morphology (Extended Data

129

Table 5 and Extended Data Fig. 2) also aligns the Irhoud specimens most closely with H.

130

sapiens, rather than with non-sapiens MP hominins and Neanderthals. They do not131

display expanded and protruding M1 hypocones, lower molar middle trigonid crests

132(especially at the EDJ), or a P4 with a transverse crest, uninterrupted by a longitudinal133

fissure. The molars are morphologically complex and reminiscent of large-toothed134

African EMH, possessing accessory features such as cusp 6, cusp 7 and protostylid on the135lower molars and cusp 5 on the upper molars. The enamel-dentine junction analysis

136

demonstrates the retention of a non-Neanderthal primitive pattern of the P4 (Extended

137

Data Fig. 2b). However, derived crown shapes shared with RMH are already expressed138

on the upper and lower molars, grouping Irhoud 11 with EMH from North Africa and the139

Levant. The lower incisor and canine roots retain a large size, but the shape is already140

within the range of the modern distribution (Extended Data Fig. 3). Mandibular molar

141

roots are cynodont, i.e. modern human-like. This mandibular root configuration of

142

Irhoud 11 is similar to that observed in EMH from Qafzeh. Finally, Irhoud 3 shows a143pattern of eruption and a period of dental development close to recent H. sapiens13.

144
145

Neurocranium

146

In contrast to their modern facial morphology, the Irhoud crania retain a rather

147

primitive overall shape of the braincase and endocast, i.e. unlike those of RMH, they are148

elongated and not globular 10,18,22. This is expressed in a low outline of the occipital

149

squama, elongated temporal bones and a low convexity of the parietal11. However, the150

frontal squama displays a rather vertical orientation and a marked convexity when151

compared to that of archaic MP specimens. These derived conditions are especially well152

expressed on Irhoud 211. A geometric morphometric analysis (Extended Data Fig. 4) of

153

external vault shape distinguishes Neanderthals and archaic MP forms with their154

primitive neurocranial shape from RMH and Upper Palaeolithic Humans. With regards

155

to PC 1, Irhoud 1 and 2 are intermediate and group together with specimens such as156

Laetoli H18 and Qafzeh as well as Upper Palaeolithic individuals from Mladeč or157

Zhoukoudian Upper Cave. To some degree all these specimens retained longer and158

lower braincase proportions compared to RMH.

159
160

The morphometric analysis of endocranial shape (Fig. 3b), which is not affected by161

cranial superstructures, shows a clear separation between H. erectus and the162

Neanderthal/archaic MP cluster along PC 2. The latter have evolved larger neocortices

163

but, in contrast to RMH, without proportional increase of the cerebellum (Extended Data

164

Fig. 5). EMH and the Irhoud hominins also display elongated endocranial profiles, but

165

are intermediate between H. erectus and the cluster of Neanderthals /archaic MP

166

hominins along PC2. They range in rough agreement with their geological age along PC1,167

in a morphological cline ending with extant globular brain shapes of RMH. Notably, Omo

168

2 falls between Irhoud 1 and 2. This similarity re-opens the question of the169

contemporaneity of Omo 1 and 223 but also raises the possibility of a late evolution of170

modern brain shape in the H. sapiens clade.

171
172

Conclusion

173

The Irhoud fossils currently represent the most securely dated evidence of the early

174

phase of Homo sapiens evolution in Africa, and they do not simply appear as

175

intermediate between African archaic MP forms and RMH. Even ca. 300-350 ka ago their176

facial morphology is almost indistinguishable from that of RMH, corroborating the177interpretation of the fragmentary specimen from Florisbad (South Africa) as a primitive

178

H. sapiens tentatively dated to 260 ka24. Anatomical mandibular and dental features, as

179

well as developmental patterns also align them with EMH. Importantly, the endocast

180

analysis suggests diverging evolutionary trajectories between early H. sapiens and MP

181

archaic African forms. This anatomical evidence and the chronological proximity

182

between the two groups25 reinforce the hypothesis of a rapid anatomical shift or even, as

183

suggested by some26, of a chronological overlap.

184
185

The Irhoud evidence supports a complex evolutionary history of our species involving

186

the whole African continent25,27. Like in the Neandertal lineage28, facial morphology was187

established early on, and evolution in the last 300 ka primarily affected the braincase.

188

This likely occurred in relation to a series of genetic changes affecting brain

189

connectivity29, organization and development22. Through accretional changes, the190

Irhoud morphology is directly evolvable into that of extant humans. Delimiting clear-cut

191

anatomical boundaries for a “modern” grade within the H. sapiens clade thus only

192

depends on gaps in the fossil record30.

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Acknowledgements:
The research program at Jebel Irhoud is jointly conducted and supported by the Moroccan
Institut National des Sciences de l’Archéologie et du Patrimoine and the Department of
Human Evolution of the Max Planck Institute for Evolutionary Anthropology. We are gratefulto the many curators and colleagues who, over the years, gave us access to innumerable
recent and fossil hominin specimens for CT-scanning or analysis, to Erik Trinkaus for providing
comparative data and to Christopher Kiarie, Mikaela Lui, Chloe Piot, David Plotzki, Alyson Reidand Heiko Temming for their technical assistance.

METHODS
Computed tomography
The original fossil specimens were scanned using a BIR ARCTIS 225/300 industrial microCT scanner, at the Max Planck Institute for Evolutionary Anthropology (MPI EVA), Leipzig,
Germany. The non-dental material was scanned with an isotropic voxel size ranging from27.4 to 91.4 µm (130 kV, 100 to 150 µA, 0.25 to 2.0 mm brass filter, 0.144 degree ofrotation step, 2 to 3 frames averaging, 360 degrees of rotation). The dental material wasscanned with an isotropic voxel size ranging from 12.8 to 32.8 µm (130 kV, 100 µA, 0.25
to 0.5 mm brass filter, 0.144 degree of rotation step, 3 frames averaging, 360 degrees ofrotation). Segmentation of the micro-CT volume was performed in Avizo (Visualization
Sciences Group). The comparative dental sample was scanned with an isotropic voxel size
ranging from 11.6 to 39.1 µm at the MPI EVA on a BIR ARCTIS 225/300 micro-CT scanner(130 to 180 kV, 100 to 150 µA, 0.25 to 2.0 mm brass filter, 0.096 to 0.144 degree ofrotation step, 2 to 4 frames averaging, 360 degrees of rotation) or on a Skyscan 1172
micro-CT scanner (100 kV, 100 µA, 0.5 mm aluminum and 0.04 mm copper filters, 0.10 to1.25 degree of rotation step, 360 degrees of rotation, 2 to 4 frames averaging). The microCT slices were filtered using a median filter followed by a mean-of-least-variance filter(each with a kernel size of three) to reduce the background noise while preserving andenhancing edges31.

Virtual Reconstruction
Using Avizo, nine reconstructions of the Jebel Irhoud 10 face were made based on
segmented surfaces of its preserved parts consisting of a left supraorbital torus, two left
maxillary fragments and a nearly complete left zygomatic bone. First, we used several
recent modern humans from diverse geographical regions (e.g., Africa, North America andAustralia) and Irhoud 1 as a reference to align the two left maxillary bones. Since a large
portion of the dental arcade of Irhoud 10 is preserved, the range of possible “anatomicallycorrect” alignments in the palate was limited (Figure 1b). Based on this maxillaryalignment, each of the subsequent reconstructions differed by several millimeters in thefollowing ways: broadening the palate; increasing the facial height; increasing the orbitalheight; or rotating the zygomatic bones anteriorly or posteriorly in a parasagittal
direction. Additionally, we aligned one reconstruction to match the facial proportions andorientation of a “classic” Neanderthal (La Ferrassie 1). In doing so, the zygomatic bonewas rotated parasagittally and moved posteriorly (> 5 mm). Correspondingly, the browridge was realigned postero-superiorly by several mm, and the maxillary bones weremoved inferiorly several mm to increase its facial height. For each reconstruction, eachbone was mirror-imaged along the mid-sagittal plane of Irhoud 1 and then the right andleft sides were merged to form one surface model. The reconstruction of the Irhoud 11
mandible was conducted by mirroring the left side of the mandible, which was best
preserved and minimally distorted, onto the right side, apart from the condyle, which was
only preserved on the right side and mirrored onto the left side. The left side of themandible was represented by three main fragments. Before mirroring, the sedimentfilling the cracks between the main fragments was virtually removed, the fragments were
re-fitted and the broken crown of the left canine was reset on its root. Note that theposition of the condyles in the reconstruction is only indicative.

Shape analysis of the face, endocast and cranial vault
Geometric morphometric methods (GMM) were used to analyse different aspects ofmorphology of the Irhoud fossils in a comparative context. To this end we digitised threedimensional landmarks and sliding semilandmarks32, 33, 34to separately analyse the shapeof the face, the endocranial profile and the external vault. On the face (Figure 3a), 3D
coordinates of anatomical landmarks, as well as curve and surface semilandmarks(n=791) were digitized using Landmark Editor35 either on CT scans (BIR ACTIS 225/300
and Toshiba Aquilion), or surface scans (Minolta Vivid 910 and Breuckmann optoTOPHE) of recent modern human and fossil crania (n =267) following Freidline et al.36.
Whenever possible, measurements were taken on scans of the original fossil; landmarks
on some fossil specimens were measured on scans of research-quality casts. Avizo wasused to extract surface files from the CT scans; data from surface scanners were preprocessed using Geomagic Studio (Geomagic Inc.) and OptoCat (Breuckmann). On theendocast (Fig. 3b), landmarks and semilandmarks (n =31) along the internal midsagittal
profile of the braincase were digitised on CT scans of the original specimens (n =86) in
Avizo (Visualization Sciences Group) following the measurement protocol described in
Neubauer et al.37, and converted to two-dimensional data by projecting them onto a leastsquares plane in Mathematica (Wolfram Research). On the external vault (Extended Data
Fig. 4), coordinate measurements of 97 anatomical landmarks and curve semilandmarks(along the external midsagittal profile from glabella to inion, the coronal and lambdoid

sutures, and along the upper margin of the supraorbital torus) were captured using a
Microscribe 3DX (Immersion Corp.) portable digitiser on recent and fossil braincases (n=296) following the measurement protocol described in Harvati et al.38. The points along
sutures were later resampled automatically in Mathematica to ensure the same
semilandmark count on every specimen.

Homo erectus samples include KNM ER 3733 (3733), KNM ER 3883 (3883), KNM WT
15000. MP archaic samples include Petralona (Petr), Arago, Sima de los Huesos H5 (SH5),
Saldanha, Kabwe, Bodo. Neanderthal samples include La Chapelle-aux-Saints 1 (LaCha),
Guattari 1 (Guatt), La Ferrassie 1 (LF 1), Forbes’ Quarry 1 (Gibr), Feldhofer (Feld), La
Quina 5 (LQ 5), Spy 1 and 2 (Sp 1, Sp 2), Amud 1 (Amud), Shanidar 1 and 5 (Shan 1, Shan
5). Primitive H. sapiens and EMH include Laetoli H18 (LH), Omo Kibish 2 (Omo 2), Singa(Si), Qafzeh 6 and 9 (Qa 6, Qa 9), Skhul 5 (Sk 5). Upper Palaeolithic modern humans
include Cro-Magnon 1 and 3 (CroM 1 CroM 3), Mladeč 1 and 5 (Mla 1, Mla 5), Brno 3,
Předmostí 3 and 4 (Pre 3, Pre 4), Abri Pataud (AbP), Cioclovina (Ci), Zhoukoudian Upper
Cave 1 and 2 (ZhUC 1, ZhUC 2). The RMH samples are composed of individuals of diverse
geographical origins (n=232 in Figure 3a, n=55 in Figure 3b, n=263 in Extended Data
Figure 4).
Crown outline analysis (Extended Data Figure 3a)
The crown outline analysis of Irhoud 10 and Irhoud 21 left M1 follows the protocol
described in Benazzi et al.39 and Bailey et al.40. For Irhoud 10, CT images were virtuallysegmented using a semiautomatic threshold-based approach in Avizo to reconstruct a 3D
digital model of the tooth, which was then imported in Rapidform XOR2 (INUS
Technology, Inc., Seoul, Korea) to compute the cervical plane. The tooth was aligned withthe cervical plane parallel to the xy-plane of the Cartesian coordinate system and rotatedaround the z-axis with the lingual side parallel to the x-axis. The crown outline
corresponds to the silhouette of the oriented crown as seen in occlusal view and projected
onto the cervical plane. For Irhoud 21, an occlusal image of the crown was taken with a
Nikon D700 digital camera and a Micro-Nikkor 60 mm lens. The tooth was oriented sothat the cervical border was perpendicular to the optical axis of the camera lens. The
image was imported in Rhino 4.0 Beta CAD environment (Robert McNeel & Associates,
Seattle, WA) and aligned to the xy-plane of the Cartesian coordinate system. The crownoutline was digitised manually using the spline function, and then oriented with the
lingual side parallel to the x-axis. Both crown outlines41 were first centeredsuperimposing the centroids of their area according to the M1 sample created by Bailey etal.40, but integrated with 10 late early and middle Pleistocene Homo M1 specimens (i.e.,
Arago-31, AT-406, ATD6-11, ATD6-69, ATD6-103, Bilzingsleben-76-530, Petralona,
Steinheim, Rabat, Thomas-3). Then, the outlines were represented by 24
pseudolandmarks obtained by equiangularly spaced radial vectors out of the centroid(the first radius is directed buccally and parallel to the y-axis of the Cartesian coordinate
system), and scaled to unit centroid size39,41. Late early and middle Pleistocene archaic
samples include Arago 31 (Ar 31), Atapuerca Gran Dolina 6-11, 6-69, 6-103 (ATD6-11,
ATD6-69, ATD6-103), Atapuerca Sima de los Huesos 406 (AT-406), Bilzingsleben-76-530(Bil76-530), Petralona (Petr), Steinheim (Stein), Rabat (Rab), Thomas 3 (Tho 3).
Neanderthal samples include Arcy-sur-Cure 39, Cova Negra, Krapina KDP 1, Krapina KDP
3, Krapina KDP 22, Krapina D101, Krapina D171, Krapina Max C, Krapina Max D, La
Ferrassie 8, La Quina H18, Le Fate XIII, Le Moustier 1, Monsempron 1953-1, Obi Rakhmat,
Petit Puymoyen, Roc de Marsal, Saint-Césaire 1. EMH include Dar es-Soltan II-NN (DSIINN), Dar es-Soltan II-H6 (DSII-H6), Qafzeh 10 (Qa 10), Qafzeh 15 (Qa 15), Skhul 1 (Skh1), Contrebandiers H7 (CT H7). Upper Palaeolithic modern humans include Abri Pataud,
Fontéchevade, Gough’s Cave (Magdalenian), Grotta del Fossellone, Kostenki 15, Lagar
Velho, Laugerie-Basse, La Madeleine, Les Rois 19, La Rois unnumbered, Mladeč 1, Mladeč
2, Peskő Barlang, St. Germain 2, St. Germain B6, St. Germain B7, Sunghir 2, Sunghir 3,
Veyrier 1. The RMH samples are composed of individuals of diverse geographical origins(n=80)
Molar and premolar enamel-dentine junction shape analysis (Extended Data Figure
3b)
Enamel and dentine tissues of lower second molars and fourth premolars weresegmented using the 3D voxel value histogram and its distribution of grey-scale values
Skinner et al. 42,43. After the segmentation the EDJ was reconstructed as a triangle-based
surface model using Avizo (using unconstrained smoothing). Small EDJ defects werecorrected digitally using the “fill holes” module of Geomagic Studio. We then used Avizo todigitise 3D landmarks and curve-semilandmarks on these EDJ surfaces42,43. For themolars, anatomical landmarks were placed on the tip of the dentine horn of the

protoconid, metaconid, entoconid, and hypoconid. For the premolars anatomical
landmarks were placed on protoconid and metaconid dentine horns. Moreover, we placeda sequence of landmarks along the marginal ridge connecting the dentine horns beginningat the top of the protoconid moving in lingual direction; the points along this ridge curve
were then later resampled to the same point count on every specimen using Mathematica.
Likewise, we digitised and resampled a curve along the cemento-enamel junction as aclosed curve starting and ending below the protoconid horn and the mesio-buccal cornerof the cervix. The resampled points along the two ridge curves were subsequently treatedas sliding curve semilandmarks and analysed using GMM together with the four
anatomical landmarks. Homo erectus includes KNM-BK 67, KNM-ER 992 (M2 and P4), S1b(M2 and P4), S5, S6a. We also included the Homo habilis44 specimen KNM-ER-1802 to
establish trait polarity. MP archaic samples include Mauer (Mauer; M2 and P4), Balanica
BH-1 (Bal). Neanderthal samples include Abri Suard S36, Combe Grenal 29, Combe Grenal
IV, Combe Grenal VIII, El Sidron 303, El Sidron 540, El Sidron 755, El Sidron 763a, Krapina
53, Krapina 54, Krapina 55, Krapina 57, Krapina 59, Krapina D1, Krapina D6, Krapina D9,
Krapina D35, Krapina D50, Krapina D80, Krapina D86, Krapina D105, Krapina D107, La
Quina H9, Le Moustier (M2 and P4), Le Regourdou (M2 and P4), Scladina (M2 and P4),
Vindija 11-39. EMH include Dar es-Soltan II H4 (DS II-H4), El Harhoura (El H; M2 and P4),
Irhoud 11 (Ir 11; M2 and P4), Irhoud 3 (Ir 3; M2 and P4), Qafzeh 9 (M2 and P4), Qafzeh 10,
Qafzeh 11 (M2 and P4), Qafzeh 15, Contrebandiers (CT; M2 and P4). The RMH samples arecomposed of individuals of diverse geographical origins (M2 sample n=8; P4 sample n=8).

Tooth root shape analysis (Extended Data Figure 3)
Dental tissues (enamel, dentine and pulp) of the anterior dentition were first segmented
semiautomatically using a region growing tool, and when possible using the watershedprinciple45; this segmentation was edited manually to correct for cracks. Each tooth wasthen virtually divided into crown and root by cutting the 3D models at the cervical plane
defined by a least square fit plane between landmarks set at the points of greatest
curvature on the labial and lingual sides of the cement-enamel junction. Following Le
Cabec et al.46 we analysed dental root shape: using Avizo, a landmark was digitised at theroot apex and a sequence of 3D landmark coordinates was recorded along the cement-

enamel junction. Using Mathematica, this curve was then resampled to 50 equidistantcurve-semilandmarks. The shape of the root surface, delimited by the cervicalsemilandmarks and the apical landmark, was quantified using 499 surfacesemilandmarks46: a mesh of 499 landmarks was digitised manually on a template
specimen, then warped to each specimen using a thin-plate spline interpolation and lofted
onto the segmented root surface by projecting to closest surface vertex. These landmarks
and semilandmarks were then analysed using GMM. Homo erectus is represented by
KNM-WT 15000 (WT 15000). The Neanderthal sample includes Krapina 53, 54, 55, 58, 59(Krp53, Krp 54, Krp 55, Krp 58, Krp 59), Saint-Césaire 1 (SC), Abri-Bourgeois-Delaunay 1(BD1), and Kebara 2 and 28 (Keb 2, KMH 28). EMH include Contrebandiers (CT), Dar esSoltan II-H4 (DSII-H4) and El Haroura (El H). Upper Palaeolithic and Mesolithic modern
samples include individuals from Oberkassel (Ob), Nahal-Oren (NO 8, NO 14), Hayonim(Ha 8, Ha 19, Ha 20), Kebara (KebA5) and Combe-Capelle (CC). The RMH sample includes
individuals of diverse geographical origins (n=47)

Statistical analysis
3D landmark and semilandmark data were analysed using GMM functions inMathematica34,47. Curves and surfaces were quantified using sliding semilandmarks basedon minimizing the thin-plate spline bending energy32 between each specimen and thesample mean shape33, 34. Missing landmarks or semilandmarks were estimated using athin-plate spline interpolation based on the sample mean shape during the sliding
process48. After sliding, all landmarks and semilandmarks were converted to shapevariables using generalised least squares Procrustes superimposition49; these data werethen analysed using principal component analysis (PCA), and between group PCA50. For
the M1 crown outlines analysis, the shape variables of the outlines were projected into theshape-space obtained from a principal component analysis (PCA) of the M1 comparative
sample. The data were processed and analysed through software routines written in R51.
Mandibular metric data (Extended Data Table 2 and Extended Data Figure 1c)
Linear measurements were taken on 3D surface models generated from microCT data in
Avizo. They were complemented by measurements of the original specimens taken by E.
Trinkaus (Extended Data Fig. 1c) and by comparative data taken from the literature52-99.
The African and European MP archaic sample includes KNM-BK 67, KNM-BK 8518, Sidi

Abderrahmane 2, Thomas Quarry I, Thomas Quarry Gh 10717, Tighenif 1, 2, 3, Arago I,
XIII, Mauer, Montmaurin 1, Sima de los Huesos XIX, XXI, XXVIII, AT 1, AT 75, AT 300, AT
605, AT 607. The Asian Neanderthals include Amud 1, Chagyrskaya 6, Kebara 2, Shanidar
1, 2 and 4 and Tabun C1. The European Neandertals include Arcy II, Banyoles, El Sidrón 1,
2 and 3, Guattari 2 and 3, Hortus 4, Krapina 57, 58 and 59, Suard S 36, Bourgeois
Delaunay 1, La Ferrassie 1, La Quina 5, La Naulette 1, Le Regourdou 1, Saint Césaire, Simade las Palomas 1, 6, 23, 59, Spy 1 and 3, Subalyuk 1, Vindija 206, 226, 231, 250, 11.39,
11.40, 11.45, Weimar-Ehringsdorf F1009 and Zafarraya. The EMH include Dar es-Soltan II
H5, El Harhoura 1, Dire Dawa, Klasies River: KRM 13400, 14695, 16424 , 21776 and41815, Qafzeh 9, 26 and 27, Skhul IV, Skhul V, Tabun C2 and Témara 1. The Upper
Palaeolithic and Epipalaeolithic sample includes individuals from Abri Pataud 1, Arene
Candide 2 and 18, Asselar, Barma del Caviglione, Chancelade, Cro Magnon 1 and 3, Dar esSoltan II H2 and H3, Dolni Věstonice 3, 13, 14, 15, 16, El Mirón, Grotte des Enfants 4,
Hayonim 8, 17, 19, 20, 25, 27, 29 and 29a, Isturitz and 115, Le Roc 1 and 2, Minat 1, Moh
Khiew, Muierii 1, Nahal Oren 6, 8, 14 and 18, Nazlet Khater 2, Oase 1, Oberkassel 1 and 2,
Ohalo II 1 and 2, Pavlov 1, Předmostí 3 and 21, Sunghir 1 and 6, Villabruna 1 andZhoukoudian Upper Cave 101, 104 and 108.

Dental metric and non-metric data (Extended Data Tables 3, 4 and 5)
Crown metric and non-metric data were collected from casts or originals with a few
exceptions taken from the literature. The latter include: Mumba XII100; Eyasi101;
Kapthurin102; Olduvai103; Sima de los Huesos103; some Sangiran metric data104. Root
metric data were taken on 3D models generated from micro-computed tomographic
data105, 106. Crown measurements were taken using Mitituyo digital calipers. Non-metric
trait expressions were scored using the Arizona State University Dental Anthropology
System107 where applicable (Lower dentition: P4 lingual cusps, Cusp 6, Cusp 7, M2 groove
pattern, protostylid; for Upper dentition: shoveling, tuberculum dentale, canine distal
accessory ridge, Cusp 5, Carabelli’s trait, parastyle, metacone and hypocone reduction),and Bailey108 for all others. RMH include individuals from South, West and East Africa,
Western and Central Europe, Northeast Asia, West Asia, India, Australia, New Guinea andAndaman Islands. For root metrics the sample composition is in Table 1 from Le Cabec etal.106. H. erectus includes individuals from Zhoukoudian, Sangiran, West Turkana, East

Rudolf, Olduvai and Dmanisi. MP African archaics (MPAf) include individuals from
Thomas Quarries, Salé, Florisbad, Rabat, Hoedijiespunt, Cave of Hearths, Olduvai,
Kapthurin, Mumba, Eyasi, Broken Hill and Sidi Abderrahmane. MP European archaics(MPE) include individuals from Mauer, Arago, Sima de los Huesos, Pontnewydd, Fontana
Ranuccio. Neanderthal samples include individuals from Amud, Arcy sur Cure, Cova
Negra, Grotta Guattari, Hortus, Kalamakia, Krapina, Kebara, Kulna, La Quina, La Fate, La
Ferrassie, Le Moustier, Melpignano, Monte Fenera, Monsempron, Montmaurin, Feldhofer,
Obi-Rakhmat, Ochoz, Pech-de-l'Azé, Petit Puymoyen, Regourdou, Roc-de-Marsal, Spy,Saint-Césaire, Subalyuk, Tabun and Vindija. EMH include individuals from Die Kelders,Equus Cave, Klasies River Mouth, Sea Harvest, Haua Fteah, Dar es-Soltan, Contrebandiers,
El Harhoura, Qafzeh, and Skhul.
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Journal of Human Evolution 15, 265-287 (1986).
Grine, F. E. & Franzen, J. L. Fossil hominid teeth from the Sangiran Dome(Java, Indonesia). Courier Forschungsinstitut Senckenberg 171, 75-103(1994).
Le Cabec, A., Gunz, P., Kupczik, K., Braga, J. & Hublin, J.-J. Anterior Tooth Root
Morphology and Size in Neanderthals: Taxonomic and Functional
Implications. J. Hum. Evol. 64, 169-193, doi:10.1016/j.jhevol. 2012.08.011(2013 ).
Turner CG, II, Nichol CR, and Scott GR. 1991. Scoring procedures for key
morphological traits of the permanent dentition: The Arizona State
University Dental Anthropology System. In: Kelley M, and Larsen C, editors.
Advances in Dental Anthropology. New York: Wiley Liss. p 13-31.
Bailey SE. 2002. Neandertal Dental Morphology: Implications for modern
human origins [Ph.D. Dissertation]. Tempe: Arizona State University. 238 p.

Figure captions

Figure 1 | Facial reconstruction of Irhoud 10, frontal (a) and basal (b) views. This Procrustes
superimposition of Irhoud 10 (beige) and Irhoud 1 (light blue) represents one possiblealignment of the facial bones of Irhoud 10. Multiple alternative reconstructions (n=9) were
included in the statistical shape analysis of the face (see Methods and Figure 3). The maxilla,
zygomatic bone and supra-orbital area on Irhoud 10 are more robust than on Irhoud 1. Scaleis 20 mm.

Figure 2 | Irhoud 11 mandibule (lateral and cranial views). See Methods for thereconstruction. The bi-condylar breadth of the Irhoud 11 mandible exactly fits the width ofthe corresponding areas on the Irhoud 2 skull. Scale is 20 mm.

Figure 3 | Comparative shape analysis. a, PCA of the facial shape. EMH and RMH are well
separated from Neanderthals and archaic MP hominins. Irhoud 1 and all nine alternative
reconstructions of Irhoud 10 (pink stars and pink 99% confidence ellipse, see Methods) fallwithin the RMH variation. b, PCA of endocranial shape. RMH (blue), Neanderthals (red) andHomo erectus (green) are separated. Archaic MP Hominins (orange) plot with Neanderthals.
Irhoud 1 and 2 (pink stars) and some EMH (black) fall outside the RMH variation. Shapedifferences are visualized in Extended Data Figure 5a. Sample compositions and abbreviations
are in Methods.

Figure 1

Figure 2

Figure 3

Extended Data Figure Captions

Extended Data Table 1| List of hominin specimens. Starting with the 2004 excavation,
specimens were given ID numbers from the project catalogue. Layer 18 of the excavation by
de Bayle des Hermens and Tixier6 corresponds to Layer 7 of the 2004-2011 excavation.

Extended Data Table 2| Measurements of the Irhoud 11 mandible after reconstruction. They
are compared to those of five groups of fossil hominins. Values in mm. x̅ = mean value, σ =standard deviation, n= sample size. The value with “?” is an estimate. Data sources and sample
compositions are in Methods.

Extended Data Table 3| Dental measurements (upper dentition) BL = Bucco-lingual width,
MD = Mesio-distal length. Values in mm. x̅ = mean value; minimum and maximum values are
between square brackets; σ = standard deviation; n= sample size. Values in parentheses
represent uncorrected measurements on worn or cracked teeth. Data sources and sample
compositions are in Methods.

Extended Data Table 4| Dental measurements (lower dentition). BL = Bucco-lingual width,
MD = Mesio-distal length, RL = Root Length. All values in mm. x̅ = mean value; minimum andmaximum values are between square brackets; σ = standard deviation; n= size of the sample.
Values in parentheses represent uncorrected measurements on worn or cracked teeth. Data
sources and sample compositions are in Methods.

Extended Data Table 5| Morphological dental trait comparison
Numbers given are trait frequencies score at the enamel surface. Sample sizes are in brackets.
Data sources and sample compositions are in Methods.

Extended Data Figure 1 | Mandibular morphology. a, Symphyseal section of Irhoud 11
mandible showing the mental angle. b, Mental area of Irhoud 11 before virtual reconstruction(top) and Irhoud 3 (bottom). Both figures are surface models generated from micro CT data.
c, Bivariate plot of mandibular corpus breadth versus height at the mental foramen. Values in

mm. Irhoud 11 plots with EMH and displays one of the largest corpus height among middle tolate Pleistocene hominins. “n” indicates sample size. Data sources and sample compositions
are in Methods.

Extended Data Figure 2 | a, shape–space PCA plot of late early and middle Pleistocene
archaic Homo, Neanderthals and RMH M1 crown outlines. The deformed mean crownoutlines in the four directions of the PCs are drawn at the extremity of each axis. Sample
compositions and abbreviations are in Methods. b, Enamel-dentine junction (EDJ)
morphology of the M2 and P4. At top left a PCA analysis of EDJ shape of the M2 places Irhoud
11 intermediate between H. erectus and RMH (along with other north Africa fossil humans)and distinct from Neanderthals. Surface models illustrate EDJ shape changes along PC1(bottom left) and PC2 (top right); the former separating H. erectus from RMH, Neanderthals
and North African EMH and the latter separating Neanderthals from RMH and north African
EMH. At bottom right a PCA analysis of EDJ shape of the P4 groups Irhoud 11 with modern andfossil humans.

Extended Data Figure 3 | Shape analysis of I2 roots. A between-group PCA shows a complete
separation between Neanderthals and a worldwide sample of recent modern humans basedon subtle shape differences. Irhoud 11 (magenta) plots at the fringes of RMH, close to the
EMH from Temara. Colour-coded Procrustes group mean shapes are plotted in the sameorientation as the I2 root surface of Irhoud 11. Although Irhoud 11 is more similar, overall, to
Neanderthals in terms of root size, its root shape is clearly modern. The Homo erectus s.l.
specimen KNM-WT 15000 and hypothetical EMH Tabun C2 have incisor root shapes similar to
Neanderthals, suggesting that roots that are labially more convex than in RMH represent a
conserved primitive condition with limited taxonomical value. Sample compositions andabbreviations are in Methods.

Extended Data Figure 4 | Shape analysis of external vault. a, Principal component (PC) scores
1 vs. 2 of external braincase shape in Homo erectus, MP archaic Homo, a geographicallydiverse RMH and Neanderthals. Results are consistent with the analysis of endocranial shape(Figure 3a). However, several EMH and Upper Palaeolithic specimens fall outside the RMHvariation. This is likely due to the projecting supraorbital tori in these specimens. b, Shapechanges associated with PC 1 (two standard deviations in either direction) shown as thin-plate
spline deformation grids in lateral and oblique view. PC 1 captures a contrast between
elongated braincases with projecting supraorbital tori (low scores in black), and a moreglobular braincase with gracile supraorbital tori (high scores in red). Sample compositions andabbreviations are in Methods.

Extended Data Figure 5 | Facial and endocranial shape differences among Homo groups.
Visualizations of GMM shape analyses in Figure 3. a, Average endocranial shape differences
Homo erectus, recent Homo sapiens, and Neanderthals. Thin-plate spline (TPS) grids are
exaggerated. b, Visualisation of shape changes along principal component (PC) 1 in Figure 3b
in frontal, lateral and superior view; two standard deviations in either direction from themean shape (grey, negative; black: positive). c, Shape changes along PC 2. All recent and fossil
modern humans (low scores along PC 2) share smaller, orthognathic faces, that differ from thelarger, robust and prognathic faces of the middle Pleistocene humans and Neanderthals (highscores along PC 1). Arrow length is colour coded (short: blue; long: red). As thesevisualisations are affected by the Procrustes superimposition, we also show TPS-grids in the
maxilla and the supraorbital area. The arrow points to the plane of the maxillary TPS (red) inthe template configuration.

Specimens

Item ID

Anatomical part

Year

Stratigraphic position

Irhoud 1

No ID

Cranium

1961

Lower deposits4

Irhoud 2

No ID

Cranium

1962

Lower deposits4

Irhoud 3

No ID

Mandible (juvenile)

1968

Lower deposits5

Irhoud 4

No ID

Humerus (juvenile)1969

Layer 18 of Tixier

Irhoud 5

No ID

Coxal (juvenile)1969

Layer 18 of de Bayle des7
Hermens & Tixier

Irhoud 6

No ID

Mandible fragment

1961-69

Identified among faunal
remains

Irhoud 7

4766

Lower right P3

2004

Initial cleaning

Irhoud 84767

Distal part of left lower
molar

2004

Initial cleaning

Irhoud 91653

Lower Molar (M1 or M2)

2006

Layer 4

Irhoud 101678, 1679, 1680, 2178,
2259

Cranium

2007

Layer 7

Irhoud 114765 + 3752

Mandible2007

Layer 7

Irhoud 12

2196

Lower incisor

2007

Layer 7

Irhoud 13

2252

Left proximal Femur

2007

Layer 7

Irhoud 14

2381, 2383

Rib

2009

Layer 7

Irhoud 15

2401

Rib

2009

Layer 7

Irhoud 16

2561, 2565

Humerus (juvenile)2009

Layer 7

Irhoud 17

2670

Right proximal Femur

2009

Layer 7

Irhoud 18

2838

Lumbar vertebra

2007

Initial cleaning

Irhoud 193747, 3748, 3749

Fibula2009

Layer 7

Irhoud 20

3751

Cervical vertebra

2009

Initial cleaning

Irhoud 21

4200

Maxilla2011

Layer A

Irhoud 224502, 4503

M2 and M3 sup2011

Layer A

Extended Data Table 1

6

Measurement

Irhoud 11

African andEuropeanarchaic MP

AsianNeanderthals

European
Neanderthals

Early modern
humans

Upper
Palaeolithic MH

Symphyseal
Height45x̅ = 31.53
σ = 3.7
n = 13x̅ = 36.1
σ = 3.36n=6x̅ = 33.98
σ = 4.64
n = 21x̅ = 36.36
σ = 6.03n=8x̅ = 31.87
σ = 2.82
n = 38Corpus Heightat Mental Foramen

38.4x̅ = 30.69
σ =4.2n = 19x̅ = 33.9
σ =3.51n=7x̅ = 31.22
σ = 3.43
n = 33x̅ = 34.23
σ = 4.57
n = 13x̅ = 30.89
σ = 3.11
n = 47

Corpus Breadthat Mental Foramen

15.4x̅ = 17.22
σ = 1.98
n = 19x̅ = 17.16
σ = 1.89n=7x̅ = 15.56
σ = 1.71
n = 33x̅ = 16.04
σ = 1.75
n = 13x̅ = 12.67
σ = 1.55
n = 48Corpus Height
at M1

36

x̅ = 31.15
σ = 4.59
n = 15x̅ =31.65
σ = 3.17n=4x̅ = 30.82
σ = 3.36
n = 22x̅ = 32.81
σ = 5.64
n = 10x̅ = 29.51
σ = 2.19
n = 29Corpus Breadth
at M1

17.7

x̅ = 17.57
σ = 2.4
n = 15x̅ = 17.54
σ = 2.67n=5x̅ = 16.7
σ = 1.75
n = 22x̅ = 17.11
σ = 2.57
n = 11x̅ = 14.25
σ = 1.57
n = 26

Corpus Heightat M1/M234x̅ = 30.82
σ = 4.21
n = 15x̅ = 32.40
σ = 1.65n=3x̅ = 29.64
σ = 3.21
n = 23x̅ = 32.88
σ = 4.26n=8x̅ = 28.64
σ = 2.3
n = 33

Corpus Breadthat M1/M2

19.3

x̅ = 18.03
σ = 2.98
n = 15x̅ = 17.57
σ = 2.25n=3x̅ = 16.35
σ = 1.56
n = 22x̅ = 17.56
σ = 2.43n=8x̅ = 14.73
σ = 1.92
n = 34

Corpus Height
at M2

31,5

x̅ = 30.42
σ =3.97n = 20x̅ = 31.75
σ = 3.82n=6x̅ = 30.10
σ = 3.4
n = 26x̅ = 32.41
σ = 5.22n=8x̅ = 27.04
σ = 2.58
n = 34

Corpus Breath
at M2

22.7

x̅ = 18.45
σ = 2.45n =20x̅ = 17.6
σ = 1.54n=7x̅ = 16.03
σ = 1.78
n = 25x̅ = 18.48
σ = 2.93n=8x̅ = 15.02
σ = 1.89
n = 36

Length of the
Dental Arcade

66.5

x̅ = 58.19
σ = 5.46
n = 11x̅ = 54.78
σ = 2.78n=4x̅ = 55.23
σ = 2.49
n = 10x̅ = 57.25
σ = 6.22n=4x̅ = 51.78
σ = 3.33
n = 26

Bigonial Breadth144 ?

x̅ = 96.93
σ = 11.84n=6x̅ = 102.13
σ = 6.22n=4x̅ = 92.57
σ = 11.62n=6x̅ = 93.75
σ = 13.93n=4x̅ = 98.59
σ = 9.67
n = 29Bicanine Breadth

38.5x̅ = 35.54
σ = 3.79
n = 11x̅ = 36.48
σ = 1.63n=6x̅ = 36.62
σ = 2.45
n = 14x̅ = 38.0
σ = 2.00n=5x̅ = 32.64
σ = 2.38
n = 28Bi-M2 Breadth

66.6x̅ = 66.13
σ = 5.81
n = 11x̅ = 72.1
σ = 1.48n=4x̅ = 69.86
σ = 3.23
n = 11x̅ = 68.46
σ = 3.54n=5x̅ = 61.75
σ = 3.88
n = 26

Bi-M3 Breadth

70.9x̅ = 70.24
σ = 6.22
n = 11x̅ = 74.86
σ = 2.78n=5x̅ = 71.9
σ = 3.29
n = 11x̅ = 72.03
σ = 4.16n=4x̅ = 66.62
σ = 4.07
n = 26

Extended Data Table 2

Irhoud 10

Irhoud21

Irhoud22

H. erectus

MPE

MPAf

Neanderthals

EMH

RMH

BL

--

9.8

--

x̅ = 10.3[9.7-11.9]
σ = 0.7 n=12x̅ =9.8[8.8-10.7]
σ =0.7 n=6x̅ = 9.7[8.9-10.5]
σ =1.1 n=2x̅ = 10.0[8.8-11.4]
σ = 0.7 n=26x̅ = 9.3[8.5-10.4]
σ = 0.6 n=11x̅ = 8.3[7.0-9.8]
σ = 0.7 n
=131

MD

--

8.9

--

x̅ = 9.6[8.5-10.3]
σ = 0.6 n=11x̅ = 8.7[7.7-9.9]σ = 0.8 n=7x̅ = 9.3[8.9-9.6]
σ = 0.5 n=2x̅ = 8.8[7.0-10.0]
σ = 0.6 n=24x̅ = 8.4[7.5-9.7]
σ = 0.6 n=10x̅ = 7.7[6.2-8.8]
σ = 0.5 n 122

BL

--

11.6

--

x̅ = 11.7[10.4-12.9]
σ = 0.9 n=12x̅ = 10.9[10.5-12.1]
σ = 1.5 n=7--x̅ =10.7[9.1-11.9]
σ = 0.8 n=30x̅ =10.4[10.0-11.1]
σ = 0.4 n=10x̅ = 9.4[7.9-11.2]
σ =0.7 n= 197

MD

--

8.4

--

x̅ = 8.3[7.4-9.1]
σ =0.5 n=14x̅ = 8.8[8.0-10.7]
σ = 0.9 n=7--x̅ = 8.0[6.2-9.3]
σ = 0.7 n=29x̅ = 7.7[7.0-8.7]
σ = 0.6 n=10x̅ = 7.1[5.6-8.6]
σ = 0.6 n=186

BL

--

11.3

--

x̅ = 11.3[9.9-13.4]
σ = 0.9 n=22x̅ = 11.1[9.9-12.2]
σ = 0.6 n=8x̅ =11.3n=1x̅ = 10.5[8.2-11.7]
σ = 0.7 n=25x̅ = 10.4[9.7-11.5]
σ = 0.6 n=11x̅ = 9.5[7.6-12.3]
σ = 0.8 n=194

MD

--

8.3

--

x̅ = 8.1[7.2-9.2]
σ = 0.6 n=21x̅ = 8.0[7.3-8.4]
σ = 0.4 n=7x̅ = 7.9n=1x̅ = 7.6[5.7-8.8]
σ = 0.9 n=25x̅ = 7.1[7.0-9.3]
σ = 0.8 n=12x̅ = 6.9[5.4-11.3]
σ = 0.8 n=173

BL

12.7

12.7

--

x̅ = 13.0[11.7-14.7]
σ = 0.9 n=18x̅ = 12.1[10.9-14.4]
σ = 1.1 n=10x̅ = 12.8[11.8-13.8]
σ = 1.0 n=3x̅ = 12.2[11.2-14.2]
σ = 0.8 n=24x̅ = 12.6[11.2-15.2]
σ = 1.3 n=22x̅ = 11.5[9.8-13.6]
σ = 0.7 n=313

MD(12.1)

12.2--

x̅ = 11.8[10.0-13.6]
σ = 0.9 n=21x̅ = 11.5[10.5-12.7]
σ = 0.8 n=10x̅ =12.4[10.0-12.4]
σ = 1.2 n=3x̅ = 11.5[8.5-13.6]
σ = 1.1 n=22x̅ = 11.5[9.9-13.9]
σ = 1.1 n=21x̅ = 10.7[8.7-13.3]
σ = 0.7 n=279

BL

12.3

12.4

13.8

x̅ = 13.3[11.3-15.5]
σ = 1.2 n=12x̅ = 13.3[11.3-16.3]
σ = 1.3 n=10x̅ = 12.5[11.2-12.2]
σ = 0.5 n=4x̅ = 12.8[11.2-16.2]
σ = 1.1 n=25x̅ = 12.3[10.5-13.7]
σ = 1.0 n=12x̅ = 11.6[9.1-14.1]
σ =1.0 n= 229

MD

10.9

11.8

12.6

x̅ = 11.6[10.2-13.6]
σ =1.2 n=10x̅ = 10.9[9.7-12.4]
σ = 0.9 n=9x̅ = 11.1[10.2-11.7]
σ = 0.8 n=3x̅ = 11.0[9.3-13.1]
σ = 1.1 n= 18x̅ = 10.3[8.6-11.8]
σ = 0.8 n=9x̅ = 9.9[7.1-12.2]
σ = 0.9 n=
199

BL

--

12.4

13.3

x̅ = 12.2[10.4-15.3]
σ = 1.4 n=11x̅ = 11.9[10.1-13.3]
σ = 1.0 n=7x̅ = 11.5[11.0-11.9]
σ = 0.6 n=2x̅ = 12.3[8.6-14.2]
σ = 1.3 n=19x̅ = 11.7[9.7-13.5]
σ = 1.0 n=8x̅ = 11.1[7.6-14.4]
σ = 1.2 n=129

MD

--

8.4

11.6

x̅ = 10.0[8.7-12.4]
σ = 1.0 n=11x̅ = 9.0[8.0-11.0]
σ = 0.9 n=7x̅ = 10.0[9.8-10.2]
σ = 0.3 n=2x̅ = 10.4[8.4-13.9]
σ = 1.1 n=19x̅ = 9.3[8.6-10.9]
σ = 0.5 n=6x̅ = 8.9[6.1-12.8]
σ = 1.1 n=115

C’

P3

P4

M1

M2

M3

Extended Data Table 3

I1

I2

C,

Irhoud3

Irhoud11

H. erectus

MPE

MPAf

Neanderthals

EMH

RMH

BL

--

6.9

x̅ =6.7[6.4-6.9]
σ =0.2 n = 3x̅ =6.9[6.4-7.5]
σ =0.4 n = 6x̅ =7.3[7.2-7.4]
σ =0.2 n = 2x̅ =7.4[6.8-8.2]
σ =0.4 n = 15x̅ = 6.6[5.8-7.6]
σ = 0.6 n = 10x̅ = 5.8[4.8-6.8]
σ = 0.4 n = 137

MD

--

(5.8)

x̅ =6.1[5.8-6.6]
σ =0.4 n = 3x̅ =5.7[4.9-7.5]
σ =0.5 n = 6

x̅ =6.2[5.9-6.4]
σ =0.4 n = 2x̅ =5.5[4.3-6.3]
σ =0.5 n = 13x̅ =5.6[4.5-6.8]
σ = 0.9 n = 10x̅ = 5.4[4.2-6.8]
σ = 0.4 n = 134

RL

--

16.1

x̅ =19.4n=1

x̅ =16.5n=1--

x̅ =17.2[13.8-20.9]
σ = 1.9 n = 17x̅ =14.4[13.7-16.6]
σ =1.2 n =5x̅ = 12.7[10.1-16.7]
σ = 1.5 n = 39

BL

--

7.6

x̅ =7.2[6.8-8.3]
σ =0.7 n = 4x̅ =7.5[6.7-8.6]
σ =0.6 n = 10x̅ =7.8n=1

x̅ =7.7[6.8-8.1]
σ =0.3 n = 17x̅ =7.1[6.4-8.0]
σ =0.6 n =10x̅ = 6.2[4.9-7.7]
σ = 0.5 n = 146

MD

--

(7.8)

x̅ =7.4[7.2-7.9]
σ =0.3 n = 3x̅ =6.6[6.3-7.8]
σ =0.4 n = 10x̅ =7.4
n=1

x̅ =6.5[5.2-7.5]
σ =0.5 n = 16x̅ =6.6[5.7-7.8]
σ =0.7 n =9x̅ = 6.0[5.0-6.8]
σ = 0.5 n = 143

RL

--

18.0

x̅ =18.6[17.1-20.1]
n=2

x̅ =16.7n=1--

x̅ =18.4[14.8-21.6]
σ =2.0 n = 15x̅ =15.4[13.7-17.3]
σ =1.6 n =6x̅ = 14.1[10.7-18.4]
σ = 1.4 n = 47

BL

9.1

9.4

x̅ =8.8[8.3-9.6]
σ =0.5 n = 5

x̅ =8.5[5.9-9.8]
σ =1.1 n = 11x̅ = 8.8[7.7-10.0]
σ =1.6 n = 2x̅ =9.2[7.8-10.3]
σ =0.7 n = 24x̅ =8.6[7.0-10.2]
σ =0.9 n =12x̅ = 7.6[6.0-9.4]
σ = 0.7 n = 132

MD

9.1

8.6

x̅ =8.4[8.0-8.9]
σ =0.4 n = 4x̅ =7.6[6.7-8.7]
σ =0.5 n = 11x̅ = 7.8[7.2-8.4]
σ =0.9 n = 2x̅ =8.0[6.9-9.0]
σ =0.5 n = 22x̅ =8.2[6.4-10.0]
σ =1.0 n =10x̅ = 6.8[5.4-8.1]
σ = 0.5 n = 124

RL

--

20.6

--

x̅ =20.8n=1--

x̅ =20.7[16.1-25.6]
σ =3.0 n = 16x̅ =17.8[16.1-19.8]
σ =1.3 n =6x̅ = 16.6[13.2-19.2]
σ = 1.8 n = 23

BL

10.2

9.6

x̅ =10.3[8.9-11.5]
σ =0.9 n = 11x̅ =9.0[8.4-10.0]
σ =0.3 n = 9x̅ = 9.7[9.4-10.0]
σ = 0.4 n = 2x̅ =9.1[7.2-10.3]
σ =0.7 n = 31x̅ =9.1[8.0-12.2]
σ = 1.2 n =10x̅ = 8.0[6.4-10.2]
σ = 0.7 n = 173

MD

9.7

9.2

x̅ =8.8[7.9-9.9]
σ =0.6 n = 11x̅ =7.9[7.4-8.4]
σ =0.3 n = 8x̅ =9.4[8.8-10.0]
σ =0.9 n = 2x̅ =8.0[6.3-9.9]
σ =0.7 n = 31x̅ =8.2[7.2-11.0]
σ =1.1 n =9x̅ = 7.1[5.8-8.6]
σ = 0.6 n = 160

BL

10.5

10.5

x̅ =10.6[9.6-11.7]
σ =0.7 n = 12x̅ =8.6[7.2-10.1]
σ =0.9 n = 11x̅ =9.9[8.7-11.1]
σ =0.9 n = 5x̅ =9.3[7.6-11.1]
σ =0.8 n = 28x̅ =9.3[7.8-10.9]
σ = 0.9 n =16x̅ = 8.4[6.8-10.8]
σ = 0.7 n = 165

MD

9.5

8.9

x̅ =8.7[7.2-9.9]
σ =0.8 n = 9x̅ = 7.4[6.6-9.5]
σ =0.8 n = 11x̅ =8.9[7.5-10.3]
σ =0.9 n = 6x̅ =7.9[5.7-11.8]
σ =1.2 n = 23x̅ =7.8[7.0-9.6]
σ = 0.9 n =12x̅ = 7.2[5.6-10.4]
σ = 0.7 n = 151

BL

12.3

12.2

x̅ =12.2[10.7-13.5]
σ =0.8 n = 15x̅ =10.6[9.7-11.5]
σ =0.6 n = 15x̅ =11.7[10.5-12.6]
σ =0.7 n = 7x̅ =11.1[9.7-12.9]
σ =0.7 n = 36x̅ =11.7[10.5-14.3]
σ = 1.0 n =19x̅ = 10.7[8.6-12.6]
σ = 0.7 n = 267

MD

14.5

12.5

x̅ =13.3[12.1-14.9]
σ =1.0 n = 13x̅ =11.2[10.6-12.0]
σ =0.5 n = 16x̅ =12.8[11.9-13.8]
σ =0.7 n = 8x̅ =11.8[10.1-13.6]
σ =0.9 n = 33x̅ =12.6[10.8-14.2]
σ = 1.0 n =20x̅ = 11.4[9.2-13.5]
σ = 0.7 n = 243

BL

12.2

12.2

x̅ =13.1[11.7-14.3]
σ =0.8 n = 14x̅ =10.4[8.6-12.4]
σ =0.9 n = 17x̅ =11.5[10.3-12.9]
σ =0.9 n = 6x̅ =11.1[9.6-12.4]
σ =0.7 n = 32x̅ =11.0[9.2-12.7]
σ = 1.0 n =22x̅ = 10.4[8.6-12.5]
σ = 0.8 n = 207

MD(15.3)

13.0

x̅ =13.3[12.5-14.4]
σ =0.6 n = 12x̅ =11.5[9.7-14.8]
σ =1.3 n = 17x̅ =12.8[12.0-13.8]
σ =0.7 n = 6x̅ =12.0[10.5-14.0]
σ =0.9 n = 29x̅ =11.7[10.2-14.2]
σ =1.1 n =15x̅ = 10.9[8.9-14.3]
σ = 0.9 n = 198

BL

--

11.1

x̅ =12.4[11.0-14.2]
σ =0.9 n = 7x̅ =10[8.7-11.3]
σ =0.8 n = 10x̅ =11.4[10.6-12.3]
σ =0.6 n = 5x̅ =10.8[7.9-13.1]
σ =0.8 n = 29x̅ =10.8[9.2-12.8]
σ = 1.0 n =14x̅ = 10.4[8.6-12.6]
σ = 0.8 n = 139

MD

--

12.8

x̅ =12.8[10.9-14.7]
σ =1.3 n = 6x̅ =11.2[9.4-12.7]
σ =0.9 n = 10x̅ =13.3[12.3-15.2]
σ =1.3 n = 4x̅ =11.8[9.4-13.9]
σ =0.9 n = 26x̅ =11.8[10.1-13.8]
σ = 1.1 n =14x̅ = 10.8[8.2-12.6]
σ = 1.0 n = 119

P3

P4

M1

M2

M3

Extended Data Table 4

Lower Dentition

Irhoud3, 11

H. erectus

MPE

MPAf

Neanderthals

EMH

RMH

P4 Lingual Cusps [pres. >1]

50 (2)

85.7 (7)50 (4)

50 (2)

97 (31)

71.4 (7)66.7 (173)

P4 Metaconid position[pres = mesial]

100 (2)90.9 (11)100 (5)66.7 (3)

97 (32)62.5(8)80.6 (177)

P4 Transverse Crest[pres. >0]0 (2)36.4 (11)40 (5)

75 (4)

90 (31)

12.5 (8)4.5 (177)

P4 Distal Accessory Ridge[pres. >0]50 (2)

75 (8)

100 (3)100 (2)87 (15)66.7 (3)47.1 (145)

P4 Mesial Accessory Ridge[pres. >0]

0 (2)

0 (7)

25 (4)

100 (2)13 (25)66.7 (3)29.5 (152)

P4 Asymmetry[pres >0]100 (2)

46.2 (13)20 (5)

95 (20)37.5 (8)0.8 (119)

P4 Fissure Pattern[pres. = U]

0 (2)

0 (5)

0 (3)

0 (2)

0 (14)

75 (4)

72.9 (145)

M1 Middle Trigonid Crest[pres. >0]0 (2)33.3 (12)88.9 (11)50 (2)

92.9 (28)35.7 (14)

1.4 (207)

M1 Protostylid [pres.>2]

50 (2)

50 (8)0 (9)33.3 (3)0 (38)

11.8 (17)0.5 (218)

M1 Cusp 6 [pres. >0]100 (2)28.6 (7)33.3 (6)100 (1)38 (21)0 (12)

18.1 (200)M1 Cusp 7 [pres. >0]100 (2)50 (12)12.5 (8)0 (2)18 (33)45.0 (20)

9.7 (236)

M2 Y Groove Pattern[pres. = Y]

50 (2)

91.7 (12)57.1 (7)0 (1)79 (33)76.9 (13)28.6 (242)

M2 Cusp number [pres.= 4]0 (2)0 (13)

0 (7)

0 (3)

0 (37)

11.8 (17)63.8 (242)

H. erectus

MPE

MPAf

Neanderthals

EMH

RMH

Upper Dentition

2

Irhoud10, 21, 22

66.7 (3)I Shoveling [pres >1]

Present (1)100 (3)--100 (1)100 (15)

83.3 (6)23.2 (122)

I2 Tuberculum dentale[pres. >1]

Present (1)

0 (2)

--

100 (1)88.9 (9)60 (5)

38.1 (118)

C Distal Accessory Ridge[pres. >0]

Present (1)66.7 (3)--100 (1)62.5 (8)100 (2)39.7 (68)

0 (2)

0 (8)

0 (4)

0 (1)

5.9 (34)

0 (10)

18.3 (243)

0 (2)

0 (8)

0 (4)

0 (1)

2.9 (34)0 (8)24.5 (241)

Present (1)0 (2)67.7 (3)100 (1)

63.6 (22)50 (10)41.2 (232)0 (2)33.3 (3)0 (1)72 (25)50 (10)46.2 (272)2

M Metacone Reduction[pres. <3.5]
2

M Hypocone Reduction[pres. <3]
1

M Cusp 5 [pres. >0]1

M Carabelli’s Trait[pres. >2]
1

M Parastyle [pres. >0]Absent (2)

0 (1)

0 (2)

0 (1)

20.8 (24)

0 (14)

0.8 (299)

M Mesial Accessory Cusps [pres>0]Absent (1)

--

0 (2)

--

45.4 (11)100 (2)

67.1 (132)1

Extended Data Table 5

Extended Data Figure 1

Extended Data Figure 2

Extended Data Figure 3

Extended Data Figure 4

Extended Data Figure 5