Modern human incursion into Neanderthal territories 54,000 years ago at Mandrin, France

Abstract

Determining the extent of overlap between modern humans and other hominins in Eurasia, such as Neanderthals and Denisovans, is fundamental to understanding the nature of their interactions and what led to the disappearance of archaic hominins. Apart from a possible sporadic pulse recorded in Greece during the Middle Pleistocene, the first settlements of modern humans in Europe have been constrained to ~45,000 to 43,000 years ago. Here, we report hominin fossils from Grotte Mandrin in France that reveal the earliest known presence of modern humans in Europe between 56,800 and 51,700 years ago. This early modern human incursion in the Rhône Valley is associated with technologies unknown in any industry of that age outside Africa or the Levant. Mandrin documents the first alternating occupation of Neanderthals and modern humans, with a modern human fossil and associated Neronian lithic industry found stratigraphically between layers containing Neanderthal remains associated with Mousterian industries.

Fig. 1.. Grotte Mandrin.

(A) Geographic location and other sites mentioned in the text. (B) Situation of the rockshelter overlooking the Rhône River Valley. (C) View looking out from the back of the shelter. AMH, anatomically modern human; MP, middle paleolithic; UP, upper paleolithic. Photo credit: Ludovic Slimak, CNRS.

Fig. 2.. Neronian points from Mandrin layer E.

Micro- and nanopoints (numbers 1 to 23), pointed micropoint (number 10), and points (numbers 24 to 33).

Fig. 3.. Mandrin layer E and layer D lithic industries.

(A) Layer D post–Neronian I Mousterian. Pseudo-Levallois points with truncated back in black exotic flints coming from ~70 to 90 km northeast of the site. (B) Mandrin layer E Neronian. Blades, bladelets, and bladelets by-products. Numbers 1 to 21, bladelets; number 18, crested bladelet; number 22, blade.

Fig. 4.. Neronian artifacts from layer E.

(A) Top to Bottom: Pointed bone point with lateral notches, worked red deer canine, eagle talon with cutmarks, and pebble with an engraved line separating the rock in two subequal parts. (B) Details of the bone point showing working traces by a flint tool. (C) Red deer canine details showing traces of scrapping by a lithic tool. These marks may have been to deliberately extract the canine or to modify its morphology.

Fig. 5.. The Grotte Mandrin human remains.

Plate illustrating the nine dental elements preserved in layers G to C (see note S5 for a detailed description of the fossil human remains). LLdm2, deciduous lower left second molar; LLM1, permanent lower left first molar; LRM3, permanent lower right third molar; URdm2, deciduous upper right second molar; b, buccal; d, distal; l, lingual; m, mesial; o, occlusal.

Fig. 6.. Geometric morphometric analyses of the crown outline and EDJ shape.

(A and B) Between-group principal components analyses (bgPCA) based on the two-dimensional (2D) landmarks Procrustes-registered shape coordinates of the crown outline of the deciduous lower second molar (Ldm2) Man11 C 204 from layer C (A) and of the deciduous upper second molars (Udm2) Man04 D 395 and Man04 D 679 from layer D (B) compared with fossil and extant hominins. (C and D) bgPCA based on the 3D landmarks Procrustes-registered shape coordinates of the enamel-dentine junction reconstructions of the Udm2 Man12 E 1300 from layer E (C) and of the permanent lower first molar (LM1) Man98 F 811 from layer F (D) compared with fossil and extant hominins. NEA, Neanderthals; UPMH, Upper Pleistocene modern humans; HH, Holocene humans; MAN_geom, geometric-based reconstruction of the Mandrin specimens; MAN_MH, modern human-based reconstruction of the Mandrin specimens; MAN_NEA, Neanderthal-based reconstruction of the Mandrin specimens (table S14).

Fig. 7.. Geometric morphometric analyses of the talon of the Udm2 EDJ.

(A) PCA based on the 2D landmarks Procrustes-registered shape coordinates of the EDJ talon shape of the Udm2 Man12 E 1300 and of the comparative fossil and extant hominin groups. (B and C) bgPCA (B) and cross-validated bgPCA (C) based on the same data. (D) PCA based on the 3D landmarks Procrustes-registered shape coordinates of the EDJ talon shape of the three reconstructions of the Udm2 Man12 E 1300 and of the comparative fossil and extant hominin groups. (E and F) bgPCA (E) and cross-validated bgPCA (F) based on the same data (table S14).

Fig. 8.. The Grotte Mandrin Bayesian model.

The model comprises the radiocarbon likelihoods and optically stimulated luminescence ages fitted within a relative age sequence that is based on the succession of archeological levels excavated at the site. A composite stratigraphy is shown at the left illustrating these stratigraphic horizons. Key probability distributions from the Bayesian model are shown on the right. These are either Boundary distributions (the top three) representing the start of a Phase, or Date ranges (the lower four) that represent the age spans of an archeological phase.

Fig. 9.. Neronian (Mandrin layer E) and Initial Upper Paleolithic points, micropoints and cores (Ksar Akil layers XXV-XX, Lebanon, Peabody Museum collections).

Drawings L. Metz. These points are precisely obtained through the same technical processes and show identical morphologies. Boxplot in upper right shows that their metrics (tip cross-sectional area; ratio width/thickness and statistic comparisons between Mandrin E points and Ksar Akil IUP points; measures at 1-mm precision) are also identical (Wilcox test: w = 4156; P = 0.1883, P > 0.01).