Studies showed that farmers themselves, rather than just their culture, expanded from the Near East around 8, cal. Paleogenetic studies also confirmed that both different Neolithization waves ultimately derived from common roots in Anatolia [ 8 , 9 ]. This spread from a common source is illustrated by the marked genetic homogeneity of early farmers, followed by a subtle genetic and temporal gradient across West Anatolia, the Carpathian Basin, Central Europe, and the Iberian Peninsula [ 8 , 10 — 14 ]. The subsequent 2, years provided evidence for relatively little genetic change, except for a mild increase of H-G ancestry in farmers, attesting to a slow but steady introgression of forager lineages into farming communities during the Late Neolithic period, in regions documented such as Central Europe and the Iberian Peninsula [ 5 , 11 ].
Nevertheless, signals of earlier admixture were detected farther west in the French territory, showing that processes of interactions between groups should have been highly variable at the regional scale [ 15 ]. In the Alsace region north-western France , after a gap of two centuries 5,—4, cal. BC , numerous cultures inherited from the LBK succeeded during the Middle Neolithic, including the Hinkelstein culture 4,—4, cal.
BC , Grossgartach culture 4,—4, cal. BC , Roessen culture 4,—4, cal. Around 4, cal. BC, the Michelsberg culture appeared, named after the archaeological site of Michelsberg in the German state of Baden-Wurtemberg [ 18 ] and developing from central Germany to the Paris Basin north of France. Simultaneously, the Munzingen culture appeared in South Alsace, mainly described through its ceramic remains.
Several hypotheses concerning the origin of the Michelsberg culture have been offered, but the one prevailing today and first introduced in [ 19 ] considers an origin in the Paris Basin [ 20 — 22 ]. This "occidental hypothesis" was reinforced by the demonstration of an important rupture in pottery traditions between the previous Alsatian cultures and the Michelsberg, combined with the continuity of specific ceramic features with southwestern traditions in the Paris Basin.
The importance and the speed of the cultural changes perceived led the author to consider that major migration should have been implied in these processes [ 20 , 22 ]. Interestingly, ceramic changes observed during the Michelsberg culture development have been accompanied by the apparition of particular funerary practices involving inhumations within circular pits. The status of these peculiar pits, as well as the status of the individuals deposited in such structures, has been the subject of intense debate.
Concerning the status of the pits, whereas C. Nickel proposed to not consider them as funerary pits but more as marginalized practices [ 27 ] , J. Lichardus and J. Schweitzer viewed them as true graves within pits especially dug for this purpose or that might have had another function in the past storage, habitation [ 28 , 29 ]. Such striking differences in funerary treatment led to the obvious question of the specific status of the individuals found in the NCV position.
Thanks to the multiplication of the discoveries and with the assistance of ethnological examples, distinct interpretations have been proposed including i graves of "relegation", i. According to the last proposal, individuals found in the NCV position can be related to "accompaniment dead" as defined by A. Testart, i. In this way, accompaniment dead could be spouses following their husbands in tombs, soldiers giving up the fight and accompanying their leader in death, subjects of a theocratic kingdom killed upon the decease of their king or, finally and most commonly in lineage societies, slaves accompanying their master in death.
According to the sacrifice hypothesis, i. A Inhumation repartition and mtDNA haplogroup distribution. Ellipses represent a circular pit; stars represent individuals in the "conventional position", whereas triangles represent individuals in the "non-conventional" position.
B MtDNA haplogroup composition of "conventional" vs. The archaeological site of Gougenheim, localized in Alsace northeast of France, 30 kilometers away from Strasbourg, Fig 2 , holds a key position in the debate surrounding both the origin of the Michelsberg groups and the significance of the mortuary treatments observed. In , excavations conducted in Gougenheim permitted the discovery of 30 circular pits containing 46 individuals, providing the most important human remains sample recovered from circular pits to date.
Radiocarbon dating conducted on human bones ranged between 4, and 3, cal. BC and permitted confirmation of the association of the large majority of the structures to the Late Neolithic and more precisely to the Michelsberg and Munzingen cultures [ 33 ].
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Located at a central point between the Paris Basin and Germany, the site thus offers a real opportunity to document the Michelsberg communities' origin. Color gradient is given according to the chronology of each group. See S6 Table for details regarding the populations considered. The archaeothanatological and biological analysis of the human remains enabled the demonstration of the combined presence of mature and immature individuals, as well as a balanced number of male and female subjects [ 33 ].
Among the 46 individuals recovered in Gougenheim, 13 were found in the CV position and 27 in the NCV position, whereas the position of the seven remaining individuals could not be described. No biological difference could be detected between the subjects found in the CV or NCV position, and no specific spatial organization of CV vs. NCV structures could be observed at the site level see Fig 1A. All evidence gathered in Gougenheim supported more probably the sacrifice hypothesis for the interpretation of mortuary practices.
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Although the ritual killing of individuals can exemplify funerary accompaniment as well as sacrifice stricto sensu , the accompaniment hypothesis seems rather implausible in Gougenheim for several reasons. Another important issue is that data compiled at Gougenheim and at the regional level suggested two separate systems of selection: an overwhelming majority of juveniles for individuals buried with the main deceased and three-quarters of adults mostly women buried in isolation [ 24 ].
Finally, several individuals had suffered a peri-mortem act of violence: parts of the body have been taken out of the pit or placed in a non-anatomical position, and in the case of individual —1, the left femur and right tibia showed burn traces. Such practices are never recounted in cases of funerary accompaniment but have been described in ritual contexts [ 24 , 36 ]. To pursue the interdisciplinary investigations developed on the exceptional site of Gougenheim, we conducted a genetic analysis of 22 subjects representative of different mortuary practices more information is provided in S1 Table.
We chose 22 individuals with exploitable teeth for DNA analyses, spatially distributed in all Gougenheim archaeological site sectors and representing all types of inhumation structures structure grouping individuals in different positions, structures grouping several CV individuals, or structure with only one NCV individual—all details about samples are present in S1 Table. While the aDNA analyses were not anticipated before the excavation, the individuals were excavated without specific aDNA care. Consequently, the teeth sampled were systematically decontaminated, i.
The samples were then reduced to powder and incubated overnight in lysis buffer 0. The procedure of Mendisco et al. This genotyping was conducted as previously described [ 37 ] to assess the DNA conservation for every sample and determine the mitochondrial maternal and Y chromosome paternal haplogroups. All primers used for these experiments and procedure details are available in Rivollat et al. Each individual's haplogroup and haplotype were determined based on PhyloTree. Sequences from all published aDNA data dating from the Mesolithic to the Late Neolithic periods were compiled 14,—2, cal.
The Gougenheim group was gathered with the data published for the Michelsberg site of Bruchsal-Aue, in western Germany [ 23 ]. The global sequence dataset was divided into 16 chronological, geographical and cultural groups in order to discuss i the implication of different anterior groups in the Michelsberg gene pool constitution and ii the relationships of the Michelsberg group with contemporaneous and later populations. Haplotype diversities S4 Table and Fst values S2 Table between populations were calculated using Arlequin software version 3. Based on these haplogroups' frequencies, PCA was plotted using R version 3.
Colors are the same as in Fig 2. We then listed all haplotypes shared between ancient Neolithic groups before 4, cal. Each remaining lineage within a given cultural dataset was traced back to its earliest match in the chronology and regarded as an ancestral lineage that arose in this culture for the first time S5 Table. From the 22 analyzed individuals, 21 exploitable mitochondrial single nucleotide polymorphisms SNP profiles could be obtained, permitting the unprecedented characterization of 21 maternal lineages haplogroups for a Michelsberg group. Among these individuals, 18 complete and authenticated HVS-I sequences or haplotypes; see the details in S1 Table could be characterized.
An exceptionally high mtDNA typing success rate Consequently, the multi-scale analyses conducted stood on the maternal gene pool characterized for the Gougenheim group. This maternal rupture could be linked to important differences in terms of genetic legacy from i Central European farmers, ii western European farmers and iii ancient European H-G.
Moreover, the ASHA permitted determining that In Germany, the resurgence of a significant number of H-G lineages was observed thereafter, around 3, cal. This genetic discontinuity appeared to be linked to new affinities with Middle Neolithic farmers from the Paris Basin, correlated to a new and important H-G legacy. All of the evidence gathered is in favor of a close genetic connection of the Michelsberg group with farmer communities from the Paris Basin region.
When chronologically and geographically replaced, the data gathered suggest that the diffusion of the Michelsberg culture was linked to a movement of human groups from West Paris Basin to East Rhine region. Since an admixture between H-G and farmers maternal components has been demonstrated in the Middle Neolithic groups of the Paris Basin Gurgy "les Noisats", 5, to 4, cal. The occidental origin in the Paris Basin of the Michelsberg culture has been proposed for a long time, based on archaeological data [ 20 — 22 ].
The major changes observed in pottery traditions between the Michelsberg culture and anterior cultures in Alsace and Germany, combined with the diffusion of specific ceramic features from southwestern traditions in the Paris Basin, led the author to consider that migrations may have been involved in such important transformations [ 20 ]. Consistently, the paleogenetic data provided by our study supported the involvement of farmer group migration to explain such a global cultural rupture. The groups involved in the Michelsberg diffusion originated in the Paris Basin and carried in Alsace few LBK-derived maternal lineages but a significant number of Paris Basin farmers and H-G maternal legacy.
The new mtDNA data gathered for the Michelsberg group also permit filling an important chronological and geographical gap to complete the scenario of H-G—farmer interactions during the Neolithic periods. Subsequently or simultaneously, such a H-G legacy appeared in the Paris Basin during the Middle Neolithic 5,—4, cal. In the meantime, very rare occurrences of H-G lineages could be observed in Central Europe, in accordance with recent genomic data [ 11 ].
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