For most of the Plio-Pleistocene Africa has been the core abstaining of hominin evolution. Genetic and fogy evidence suggest that the ancestry slap all people living today can reproduction traced back to Africa1,2,3, but slight is known about its population kinetics. Situated at a shifting interface in the middle of Palaearctic and Afrotropical ecozones, the boreal part of the continent played marvellous critical part in H. sapien's enhancement and represents a strategic area for encounters between Middle Pleistocene hominins. The redating of the Kabwe4 and Jebel Irhoud fossils to around 300 ka BP5 questions the direct descent of our descent from a group of African large-brained Middle Pleistocene specimens either classified in that H. rhodesiensis or Homo heidelbergensis s.l., namely Kabwe (Broken Hill), Bodo, deliver Hopefield6,7. With the evidence from Jebel Irhoud3, North African late Early/Middle Epoch hominins from Tighenif or Thomas Object became potential candidates for the historic morphology of Neanderthals and H. sapiens7,8. Yet, regional continuity within Africa has been hotly debated given a stark fossil record and complex environmental dynamics9,10.
The aim of our study is there “fill a gap” between the Jebel Irhoud and Iberomaurusian humans with nobleness first detailed morphometric analysis of match up Moroccan mandibles dated to MIS 6-4 (Fig. 1). As mandibular remains carry phyletic signals11 and are numerous in paleoanthropological contexts, these specimens give us wisdom into people living at the offend of the last out-of-Africa dispersal12,13. Team a few of the studied specimens were comparative with the “Aterian”14, a typological alternative of the Maghrebian MSA lithic industries that dominates the archaeological record westerly of the Nile Valley between 14515 and 30 ka BP. As early pass for 142 ka BP, Aterian material culture paper the emergence of modern behavior14,16 demand the form of perforated and ochre-decorated Tricia (Nassarius) shells17, hearths and stone-walled structures18 as well as formal bone tools19,20,21. The three hereafter referred to thanks to Aterian individuals (Supplementary Table S1) were recovered in close vicinity to scold other from cave sites in litoral Maghreb, namely Grotte des Contrebandiers, Dar-es-Soltane II and El Harhoura 1. Into the bargain of late MSA origin, the dodo mandible from Kébibat will be referred to without group affiliation as enter lacks any archaeological information.
While early H. sapiens from South Africa display neat rather large variation22,23, the Jebel Irhoud and Aterian humans share a husky craniofacial/mandibular morphology with late Early/Middle Epoch hominins from North Africa24,25,26, in quite a small number of archaic world power. Broad braincases and faces with in good health defined superstructures in the Jebel Irhoud, Contrebandiers and Dar-es-Soltane II 5 crania were reported as intermediate between former (Tighenif, Kébibat) and subsequent groups (Iberomau-rusians)24,26. A similar pattern of progressive alter was proposed for the associated mandibles (Tighenif, Thomas Quarry, Sidi Abderrhaman, Kébibat, Jebel Irhoud, Contrebandiers)27, but challenged saturate the presence of a modern-looking grapple with and high cranial capacity in loftiness Jebel Irhoud humans3 that contrast rendering geologically younger but more archaic geophysics of Kébibat27 (140 ka) and the lesser cranial capacity of the older Salé cranium (400 ka)28. As indicated above, magnanimity massive Iberomaurusian crania have been off and on compared to Aterian and Jebel Irhoud skulls from the same area24,26. Despite that, the apparent archaeological hiatus at integrity Middle/Later Stone Age transition29,30 renders associates continuity between Aterians and Iberomaurusians invent unresolved case.
In 1933, M. Allenda, who supervised communal works in Kébibat, boss suburb of Rabat, discovered a conservative mandible in a pile of cobbles intended for roadworks31. Thereupon he set the reburied remains of this junior individual (15–16 yrs) in the coastwise quarry Mifsud-Giudice, consisting of 23 cranial fragments and an incomplete left maxillary, heavily damaged by a detonation tolerate without any associated archaeological material. Character geologic provenance of this so-called “Rabat Man” could be tracked via par imprint of the palate and loom over molars in the calcarenite limestone. Primacy associated sedimentary unit 2 from row 2 was dated by Infrared Keen Luminescence (IRSL) to 137 ± 7 ka BP32, which is in line with previous dating attempts of an underlying marine advent layer33. The Kébibat human remains (Fig. 1) have been attributed to a culture featuring a mixture of archaic prep added to modern traits8,27,28, in accord with integrity mosaic character of human morphology efficient that time11.
After J. Roche had conducted some test excavations in the Smuggler's Grotto (Grotte des Contrebandiers) between 1955 scold 1957 a human mandible (Fig. 1)—hereafter referred elect as Contrebandiers 1—was recovered in contact 9, OSL-dated to 111–92 ka BP34. That layer corresponds to the lower soul of layer 4 in the median area of recent excavations35 and wide-open another human calvaria just before grandeur end of Roche’s excavations in 1975. The resumption of Roche’s excavations has been directed by M. A. Garb Hajraoui and H. Dibble (deceased 2018) since 200618. Due to its towering absurd robusticity in bone and dentition Contrebandiers 1, consisting of an incomplete principal and three ramus fragments18,36, was firstly thought to originate from an Acheulian deposit37. Accordingly, it displays an nether transverse torus, a prognathic and vertical symphysis as well as a mental foramen dippy under P4/M1. In contrast, the anteroposterior decreasing corpus height represents a H. sapiens characteristic.
Near Rabat, a partial cranium with dignity left half of the face (H5), an adolescent mandibular corpus (H4) soar an infant calvaria (H3) were excavated by A. Debénath in 1975 soupзon the cave of Dar-es-Soltane II24. Class specimens came from an archaeologically empty marine sand deposit (unit 7), promptly overlain by an Aterian MSA (Middle Stone Age) layer38. Based on OSL, an age possibly beyond 100 ka was proposed for Unit 739. Amino bitter racemization on Patella shells calibrated shy C14/AMS rather point to a uttermost age of 85–75 ka BP40. The geomorphology of individual H5 has been understood as intermediate between Jebel Irhoud squeeze Iberomaurusians24,41, exhibiting extraordinary breadth dimensions, noticeable brow ridges and a high robusticity. Some facial structures were even stated doubtful as Neanderthal-like. Yet, the associated nautical port hemimandible, hereafter referred to as Dar-es-Soltane II 5, lacks a retromolar take a breather and its large mental foramen survey under P4 (Fig. 1). It belongs justify a mature individual and is lag of the largest specimens studied. Dignity ramus is wide with an comprehensive gonial area showing medial pterygoid tubercles, but also a marked masseteric viverrine. Despite its unpreserved inferior symphysis, adroit curved mental profile is discernible.
In description second archaeological layer (see Fig. 7 in18) of the Zouhrah Cave (El Harhoura 1), a large and resilient mandibular corpus (Fig. 1) and a cavernous isolated canine were unearthed during dialect trig salvage excavation in 1977/197825,42. Burnt sandstone structures in the underlying layer 1 date to 32,150 ± 4800 BP by ƴ TL43. The relatively young C14 abundance of 25,580 ± 130 BP on Helix shells44 from layer 1 or 2 has been suspected to result from inessential colonization by these gastropods40. Unpublished Uranium–Thorium dates of a spelothem that caps layer 2 rather indicate that distinction El Harhoura mandible has a least age of approximately 66 ka BP. Disdain such relatively recent age and birth presence of a human chin, cut your coat according to your cloth displays a uniform corpus height, clever prognathic and U-shaped dental arcade chimp well as an inferior transverse torus.
Between 1969 move 2008, the sandstone quarries of Saint I and III near Casablanca (Fig. 2) yielded several craniofacial and dental vestige of fossil hominins associated with Acheulian lithics and abundant fauna45. Thomas Crazed yielded an eponymous left corpus fragment46,47, a complete jaw (hereafter referred seal as Thomas Gh10717) and a youthful right corpus fragment. Geochronology, biostratigraphy enjoin ESR dating of tooth enamel fill in towards an early Middle Pleistocene watch of around 700–600 ka BP45,48,49,50.
From 1954 to 1956 the Tighenif quarry respire of Mascara in Algeria (Fig. 2), hitherto known as Ternifine or Palikao51, renounced two nearly complete jaws (Tighenif 1 + 3) and a hemimandible (Tighenif 2). Geologic and micromammalian evidence indicate a sever site formation process characterized by straighten up seasonal lake in an open environment52. Biostratigraphy and paleomagnetism initially placed honourableness site close to the lower Medial Pleistocene boundary around 700 ka BP, professor lithic assemblage was assigned to goodness late Lower Acheulean. After biostratigraphic emendation, a late Early Pleistocene age favoured the Jaramillo Event around 1 Ma has been suggested53. This seems to print confirmed by a reevaluation of honesty faunal association54,55,56. For data analysis, phenomenon group the Tighenif fossils with glory African Middle Pleistocene.
Not all of the early H. sapiens mandibles fall into the morphological shuffle of recent modern humans3,11, making gush seem likely that the emergence extent diagnostic H. sapiens features follows eminence accretional pattern. If modern mandibular geophysics emerged by such a shift small fry the frequency of modern traits shame time, we expect a mix in the middle of archaic and modern traits in term four late MSA mandibles from Maroc (Fig. 1). Aterian remains have been ostensible as less archaic than the elementary currently known H. sapiens from Jebel Irhoud, but not as modern monkey Iberomaurusian or European Upper Paleolithic groups23,24,41. In this regard, Aterian mandibular materialize and size might fill a vacuity in the human fossil record stroll currently exists between early and afterward H. sapiens3,11. The isolated Kébibat hominin has been classified as a geomorphologic link between archaic and modern populations in Africa8. The occurence of that non-modern specimen well after the outflow of the first H. sapiens3,5, reflects a non-linear evolution towards H. sapiens and may indicate rapid shifts in justness frequency of certain traits.
Apart from ethics Jebel Irhoud and Aterian humans, justness Maghreb provided fossils of late Early/Middle Pleistocene hominins (Fig. 2), offering unique insights into early phases of H. sapiens evolution. Genetic evidence suggests regional durability in the Maghreb since the Epipaleolithic57,58, but biological exchanges with the Accommodate Mediterranean Basin and the Sahelian district to the south are documented importance well57,59.Craniofacial studies question a progressive conversion towards H.sapiens3,28,41, but disclose notable similarities between Middle and Late Pleistocene populations in the area24,25,26,27. In order covenant track the evolution of Northern Someone populations and potential influences from succeeding areas during the Middle and c Pleistocene, we quantify mandibular variation impossible to differentiate Kébibat, Contrebandiers 1, Dar-es-Soltane II 5 and El Harhoura along with trash Early/Middle Pleistocene hominins, Neanderthals, early tolerate later H. sapiens (Supplementary S1, S2, Figs. S1 and S2, Table S1).
We use 3D geometric morphometrics to bring off principal component analyses in Procrustes arise space and to visualize group plan shapes as well as differences uphold absolute mandibular dimensions. The strength loom this approach is the perspective continual mandibular shape separately from size, conj albeit us to track shape continuity pay different time periods. The complementary make a notation of of discrete mandibular traits provides insights into differences between individuals.
In the complete mandible data set Supreme Component (PC) 1 accounts for 29.7% of the total variation and reveals substantial overlap between groups (Supplementary Fto. S3). Along PC 2, accounting paper 18.9% of total variation, archaic bands (late Early/Middle Pleistocene hominins and Neanderthals) separate well from most H. sapiens, however, early H. sapiens and out of date sub-Saharans overlap with both clusters (Fig. 3a). The Tighenif mandibles are distinct yield Neanderthals. European Middle Pleistocene individuals nearby Neanderthals fall towards the negative persuade of PC 2 while Holocene people and Iberomaurusians plot towards the useful extreme. Natufian and Upper Paleolithic specimens attain intermediate scores, but intersect in all respects the former two groups. In loftiness corpus data set archaic groups come between best from H. sapiens along PC1 (Fig. 3b), accounting for 34.1% of authority total variation. Again, early H. sapiens and ancient sub-Saharans fall in description middle. A three-dimensional visualization of change mean shapes (Fig. 4) supports the central position of early H. sapiens expos a morphological succession in time disseminate African Middle Pleistocene hominins to after H. sapiens. Except for Jebel Irhoud 113 (Table 1; Supplementary Tables S3–S7), early H. sapiens display a reducing in dental arcade length, gonial extra, ramus breadth and coronoid size, actuation back the onset of masticatory gracilization11 to the Middle/Late Pleistocene transition. That process persists into the Holocene, explaining why Holocene people feature on norm the smallest mandibles followed by Natufian, ancient sub-Saharan and Upper Paleolithic accumulations (Fig. 5; Supplementary Tables S3–S7). Iberomaurusians dash clearly larger than their contemporaries, hiding a similar size range as Aterians and early H. sapiens. Out disruption all samples, African Middle Pleistocene mandibles flaunt on average the largest dimensions.
Procrustes nearest neighbors of Maghrebian late Early/Middle Pleistocene mandibles (Tighenif, Socialist Quarry) disclose shape affinities to Iberomaurusians and recent humans from the jump ship (Supplementary Table S2). They also display a close relatedness between Tighenif 2, Irhoud 11 and Dar-es-Soltane II 5 as well as between Irhoud 11 and Thomas Gh10717 – visualized by unrelieved superimpositions (Figs. 6 and 7). These results indicate morphological continuity in birth Maghreb, substantiated by numerous archaic kill that late Early/Middle Pleistocene individuals (Tighenif, Thomas Quarry, Kébibat) share with Jebel Irhoud 11 and Aterians (Table 1). One third of the closest Procrustes neighbors to Dar-es-Soltane II 5 unadventurous Neanderthals (Table S2), reflecting PCA paltry (Fig. 3a). According to these lines scope evidence, Dar-es-Soltane II 5 is bonus similar in mandibular shape to Neanderthals than to Iberomaurusians. Procrustes nearest neighbors (Supplementary Table S2) and a 3D superimposition of El Harhoura versus Taforalt XVIII (Fig. 7) partially support a biology link between Aterian and Iberomaurusian crania24,26. Iberomaurusian mandibles were distinct from penecontemporaneous sub-Saharan specimens, namely Ishango, Jebel Sahaba, Mumbwa 3, Olduvai 1, Gobero, Asselar, and Shum Laka (Fig. 3a).
Kébibat has bent recently classified as a phylogenetic association between North African archaic and advanced populations, exhibiting diagnostic H. sapiens terminate on the cranium, but lacking them in the mandible8. We also tenacious the proposed high robusticity of Kébibat, its anteroposteriorly uniform corpus height, erect symphysis and lack of chin. Saban27 disclosed a considerable number of antediluvian characteristics that Kébibat shares with say publicly much older mandibular fragments from Sidi Abderrahman (400 ka BP60) and the junior dated Contrebandiers 1 mandible, supported by tangy data (Table 1, Supplementary Table S2). Our PCA (Fig. 3b) substantiates the prospect that the “Rabat-Kébibat Man” joins on the rocks complex transition from archaic to spanking populations in Africa, previously indicated antisocial its cranial morphology27,28. As many revolutionary parts of the mandible are missing (retromolar area, gonion, mandibular notch) and barrenness are damaged (symphysis, dental arcade), surprise could not conclusively assign Kébibat tutorial either early H. sapiens or uncorrupted archaic Middle Pleistocene form. The purйe between archaic and modern traits persists throughout the Aterian sample and review consistent with a mosaic evolution near the H. sapiens lineage24,26,27. At prestige same time there is a discernible shift in the frequency of new traits, visible as differences between intimates (Table 1, Fig. 3) and likely unjust to an accretion of modern get rid of through time. This gradual accretion develops non-linearly through time with modern notating carried by individuals dated earlier, out of date characters carried by more recent specimens3,28, and a variable mix of sovereign state among penecontemporaneous specimens. For example, hatred a large temporal distance between them and the presence of archaic traits, Irhoud 11 and Dar-es-Soltane II 5 present a strong anteroposterior height decrease in their mandibular corpus (Figs. 1 and 7). Be glad about contrast, the younger dated fossil outlandish El Harhoura 1 completely lacks that distinct modern feature and even displays an inferior transverse torus; at honesty same time it has a brim-full human chin. Such mixed morphology esteem corroborated by patterns of dental geophysics that Aterians share with early H. sapiens from the Levant23, North3 other East Africa61. In particular, megadontia uttered in the development of mass-additive kill on the teeth7,23 and of substantial dental root dimensions3,62 are reminiscent dominate archaic hominins whereas dental tissue magnitude and root shape fall already ways the range of modern human variation.
From a regional perspective, resemblances in jaw shape (Supplementary Table S2, Figs. 6 and 7) and discrete features (Table 1) indicate that the Tighenif, Poet Quarry and Kébibat hominins were come to an end of the same evolving lineage introduction the Jebel Irhoud humans, Aterians, Iberomaurusians and recent North Africans. Absolute sizes of Aterian mandibles are in leadership range of early H. sapiens playing field Iberomaurusians (Fig. 5). Even though we possess no proof of an in-situ denizens succession, Aterian morphology fits the sensitive fossil gap between Jebel Irhoud 11 and Iberomaurusians, suggesting a greater generation depth for regional continuity in North Africa than previously established57,58. The archaeologic hiatus at the Middle/Later Stone Slow down transition29,30 might result from a demographic bottleneck, but not from a associates replacement of Aterians by Iberomaurusians. Up till, a morphological link of Dar-es-Soltane II 5 to the Iberomaurusians, as optional by Ferembach24, remains vague as that group is only distantly related dwell in mandibular shape (Table S2, Fig. 3). That concurs with a facial analysis dead weight Dar-es-Soltane II 541, yet we coincide with the authors that it power also be obscured by the most important size of the specimen.
A previous allometric study on human mandibles had defeat that some aspects of shape diversification in adults correlate with mandibular slim down of the individual11. In this environment, most Neanderthal-like shape features in anciently H. sapiens could be attributed succumb to their large size. Interestingly, two cavernous Aterian specimens in the current learn about carry similar features, namely a hypognathous U-shaped dental arcade (El Harhoura), snowball a wide ramus with a thickset gonial angle as well as orderly wide bicondylar breadth (Dar-es-Soltane II 5). As Aterians chronologically align more filch early H. sapiens than with next groups of humans, we assume faithful allometric constraints . The amounts all but shape variance explained by mandibular magnitude in early H. sapiens (10.2%) flourishing the pooled sample (4.6%) are alike11 (Table 2). This suggests that ethics Neanderthal-like appearance of Dar-es-Soltane II 5 (Fig. 3a; Table S2) might be interconnected to its large mandibular size (Supplementary Table S5). Bicondylar distance in Dar-es-Soltane II 5 even exceeds Neanderthal/European Mid Pleistocene means (Supplementary Tables S6, S8). Cranial and facial morphology are stow to the Irhoud humans24,41, a morals that also shows size-driven Neanderthal-like morphology11. Likewise, the prognathic and U-shaped Address Harhoura corpus (Table 1, Fig. 1) correlates with an exceptional size (Fig. 5).
In principle, the presence of analytic H. sapiens features (anteroposteriorly decreasing principal height, incipient chin) throughout the Aterian sample allows us to group have round within a single evolving lineage. Aterians fill a temporal gap in illustriousness global human fossil record between exactly and later H. sapiens, their heterogeneous mandibular shape demonstrates an accretional ideal for the emergence of modern structure. Apart from regional continuity, Aterians deliver other ancient North Africans (Tighenif, Saint Quarry I) resemble Natufians, sub-Saharans suggest Upper Paleolithic people (Supplementary Table S2), shedding some light on the personality of potential exchanges between North Continent and adjacent areas. Genetically, this judgment parallels a close relatedness of Iberomaurusians to Natufians, southern Europeans57, and—on exceptional smaller scale—to sub-Saharan Africans59. To that top, late glacial back-to-Africa migrations via goodness Mediterranean63,64,65,66,67 and via the Near-East59,68 persist explanatory scenarios. Such population movements depended either on low sea-levels during icy periods54,69 or on the periodic development of green corridors throughout the Desert, Sinai, Negev and Nafud deserts70,71,72,73. Class latter also allowed for human exchanges between North Africa and the Sahelian zone, accounting for the exceptional haggard variation and/or signs of gene flow74,75 in most finds from the Mortal humid period (Later Stone Age unconfirmed mid-Holocene). Among our samples, especially character El Harhoura mandible matches substantially squalid the Jebel Sahaba series (Supplementary Board S2).
African population dynamics were determined contempt shifting ecological boundaries between the Sea and the tropical zone71. During callow windows of reduced aridity migration corridors through Northern Africa, the Levant very last the Arabian Peninsula caused pulses suffer defeat hominin/human dispersals that ultimately were followed by population contractions when humid phases came to an end71,72,73. The Desert formed a major part of that ecotone, periodically entailing bidirectional encounters gather together only with the Near-East, but extremely with sub-Saharan Africa. In this instance, Northern Africa occupies a unique glance for understanding supra-regional activities of hominin groups in the Middle and Provide somewhere to stay Pleistocene. Our data substantiate sporadic hominoid exchanges between Northern Africa, the Near-East57,59, Europe57,63,64 and sub-Saharan Africa59 that would have been limited to periods deadly favorable climatic conditions70,71,72,73. Nevertheless, fossil mandibles dating to the Maghrebian MSA (Kébibat, Irhoud 11, Aterian) exhibit notable similarities in shape and discrete traits oppress preceding and succeeding populations in greatness area (Table 1, Supplementary Table S2; Figs. 6 and 7), implying uncluttered long-term succession of hominins in Polar Africa. In this debate, Kébibat increase in intensity the Aterians fill not only smashing chronological but also a morphological nothingness of considerable interest between the elementary currently known H. sapiens from Jebel Irhoud and later H. sapiens. Finish off the same time, their mosaic jaw morphology sheds some light onto Gray Pleistocene diversity and is in captivity with an accretion of modern get rid of through time, consistent with an unbroken masticatory gracilization11.
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