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Post by Admin on Oct 15, 2019 19:39:31 GMT
Investigations of the genetic relationships among humans from multiple Neolithic sites across western Eurasia have shown that Neolithic lifeways dispersed across Europe via a large-scale process of migration (1⇓⇓⇓⇓–6) starting from Anatolia and the areas of the Aegean at ca. 7000–6500 (cal) BCE (7⇓⇓–10). In Europe, migrating people and Neolithic lifeways dispersed along two main routes: an inland route (partly along the Danube River) and a route along Mediterranean coastal areas (11⇓–13). Around 4000 BCE, Neolithic farming communities reached the northwestern fringes of Europe, including the British Isles (14, 15) and Scandinavia (1, 2, 16, 17). A marked hunter-gatherer (HG) admixture has been observed in the later farmer groups compared with the Early Neolithic farmers on the continent (2, 10, 12). During this period of important social and demographic change, a new phenomenon of constructing megalithic monuments emerged, starting around 4500 BCE in France (18), 3700 BCE in the British Isles (14, 19⇓⇓⇓⇓⇓⇓–26), and 3600 in Scandinavia (16, 27). These Neolithic megalithic tombs are concentrated along the Atlantic coastal areas, stretching from the Mediterranean to Scandinavia, including the British Isles and regions in the northern European plain (28), but also in southern France, northern Italy, and on the Islands of Corsica and Sardinia (Fig. 1) (19, 27). Fig. 1. Map of Europe with megalithic burial sites (red squares) and nonmegalithic sites (red circles) from this study, and comparative published data from megalithic sites (black squares) sequenced to date in Europe (Dataset S1.3). The date range represents the 95% CI of available samples from these sites, except for La Mina in Spain. Blue shading represents the estimated distribution of early megalithic burials. Bold italic type indicates dates (95% CI) estimated for the start of dolmens and passage grave monuments, based on samples from these contexts. Regular text indicates time interval associated with the earliest cultural material in the megaliths (27, 45). The emergence of these megaliths was closely associated with the development of farming communities (14, 23, 25, 27, 29), but the origin and the social structure of the groups are largely unknown. The similarities in the construction and design of some types of megaliths (i.e., dolmens and passage graves) from Iberia to southern Scandinavia, Britain, and Ireland is compelling. Interregional interaction has been evidenced in the same period from the dispersal of domesticated resources, raw materials, and artifacts, possibly reflecting shared social and cultural systems as well as shared cosmology of the groups (21, 27, 28, 30). Although it is clear that many megaliths were used for collective burials (27, 29, 31), it has been difficult to evaluate which members of the communities were buried in the tombs. Some assemblages include males, females, juveniles, and children, implying familial burials. Many tombs have poorly preserved human remains and also show secondary usage in later times, complicating assessments. The use of megaliths as burial grounds for the community as a whole would imply some level of shared ideology over vast geographical areas (31, 32). However, it has also been argued that the monumental burials and associated rich material culture reflect the emergence of social differentiation or stratification (33⇓⇓–36; see ref. 37 on segmentally structured societies), with the monuments perhaps symbolizing status and territorial markers (37⇓⇓–40). Table 1. Summary of genetic and archaeological information about the 27 individuals in the study
Radiocarbon date (95% CI, cal BCE) Sequence coverage Haplogroup Estimated contamination Individual Site Upper Lower nuDNA mtDNA Sex mt Ychr mtDNA 95% CI Autosomal Primrose 2 Primrose 3790 3660 5.76 817.93 XX H1+16189 0.05 0.01–1.22 1.283 Primrose 17 Primrose 3780 3650 0.19 49.51 XY K1a+195 I 0.66 0.11–21.63 0.049 Primrose 18 Primrose 3770 3650 0.10 55.71 XY K1a+195 I 0.59 0.10–18.30 0.000 Primrose 12 Primrose 3770 3650 0.25 325.42 XY W1+119 I2a2a1a1a2 0.09 0.01–2.62 0.000 Primrose 3 Primrose 3770 3650 0.22 125.69 XY H1i I 5.28 1.91–12.50 0.000 Primrose 16 Primrose 3690 3530 6.40 442.67 XY K1a4a1 I2a2a1a1a 0.06 0.01–1.53 0.951 Primrose 10 Primrose 3640 3520 0.23 178.60 XY K1a+195 I 0.17 0.03–5.23 0.000 Primrose 6 Primrose 3640 3380 0.27 1,158.06 XX K1a+195 0.03 0.00–0.84 0.000 Primrose 13 Primrose 3630 3370 4.73 675.01 XY T2b3c I2a2a1a1a 0.03 0.01–0.64 1.731 Primrose 7 Primrose 3510 3360 0.01 43.44 XY K1a4a1 NA 1.44 0.18–14.26 0.000 Primrose 9 Primrose 3500 3360 7.10 923.93 XY U5b2c I2a2a1a1a 0.03 0.00–0.88 1.520 Carrowmore 4 Carrowmore 3640 3380 0.04 451.69 XY T2c1d1 I 0.03 0.00–0.72 0.100 Midhowe 1 Midhowe 3630 3370 0.27 22.00 XY H5+16311 I2a1b 1.52 0.24–44.17 1.150 Lairo 1 Lairo 3360 3100 0.22 25.08 XY U5b2 I2a1b 0.96 0.16–31.07 0.022 Balintore 4 Balintore 3370 3110 1.54 168.43 XX H1 0.18 0.03–4.71 0.033 Midhowe 2 Midhowe 3360 3100 0.25 29.38 XY K1a+195 I 0.75 0.14–23.16 0.281 Ansarve 5 Ansarve 3500a 3130* 0.13 114.73 XX K1a2b* 0.21 0.04–7.79 0.000 Ansarve 3 Ansarve 3490a 3110* 0.14 300.87 XX T2b8* 0.04 0.01–1.02 0.046 Ansarve 8 Ansarve 3340a 3030* 1.94 1,462.38 XY J1c5* I2a1b1a1† 0.01 0.00–0.14 0.441 Ansarve 14 Ansarve 3330a 2950* 2.58 431.47 XY J1c5* I2a1b1a1† 0.02 0.00–0.41 0.525 Ansarve 17 Ansarve 3330a 2930* 6.80 491.04 XY HV0a* I2a1b1a1† 0.06 0.01–2.06 1.461 Ansarve 6 Ansarve 3090a 2920* 0.0027 137.06 XY J1c8a* NA 0.06 0.01–1.70 NA Ansarve 7 Ansarve 3010a 2890* 0.0014 24.54 XY K2b1a* NA 0.33 0.06–8.90 NA Ansarve 9 Ansarve 2880a 2630* 0.0009 26.73 XX K2b1a* 0.29 0.05–6.99 NA Ansarve 16 Ansarve 2810a 2580* 0.33 23.17 XY H7d* I2a1b† 1.60 0.27–46.97 0.004 Kolin6 Kolin 4910 4740 1.51 218.40 XX H+16129 0.10 0.02–2.23 2.639 Kolin2 Kolin 4650 4460 0.10 42.39 XX W1+119 0.37 0.06–10.83 0.068
Some scholars hypothesize that the people buried in the megalithic structures were kin related (41⇓–43). Analyses of mitochondrial data (mtDNA) from megalithic burials at Falbygden and Gotland in modern-day Sweden have revealed a large lineage variation, and thus the groups did not seem to have been organized matrilineally (44, 45; however, contra ref. 43). Genomic data are necessary to provide deeper information on kin relations and the social dynamics and general social structure of the societies or groups. However, as genomic data have been available from only a few individuals from megalithic burials, the origin and dispersal dynamics of the funerary practices, as well as the population history of the people that used the burial constructions, have also remained uncertain. In the present study, we investigated the genetic structure and demographic affinities of people buried within megaliths to shed light on this burial phenomenon, the social dynamics of the people buried in the monuments, and their demographic history. We generated and examined genome sequence data from 24 individuals from five megalith burial sites located in Ireland, the Orkney Isles, and the Island of Gotland in the Baltic Sea dated between ca. 3800 and 2600 cal BCE encompassing wide-ranging examples from the megalithic tradition in northern Europe. The study also incorporated three individuals from nonmegalith contexts from mainland Scotland (3370–3100 cal BCE) and the Czech Republic (4825–4555 cal BCE) (Table 1).
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Post by Admin on Oct 15, 2019 22:13:04 GMT
Results We present genome data from 27 individuals excavated from European Neolithic contexts, of whom 24 were buried in megaliths; Primrose Grange (n = 11) and Carrowmore (n = 1) in Ireland; Lairo (n = 1) and Midhowe (n = 2) in the Orkney Islands, Scotland; and Ansarve (n = 9) in the island of Gotland, Sweden (16, 45, 46) (Table 1 and SI Appendix, section S2). Individuals from the Scottish “short cist” burial Balintore (n = 1) and the Czech Republic Kolin Rondel site (n = 2) (46), associated with the Stroked Pottery culture, were also investigated. These individuals were all radiocarbon-dated to between 4825 and 2580 cal BCE (Table 1). We compared our data with genetic data previously generated from 36 individuals from 16 megalithic sites (Fig. 1 and Dataset S1.3), as well as with farmer groups of nonmegalithic contexts (Dataset S1.3), to investigate the population history of people buried in megaliths. The individuals buried in these megaliths from the British Isles and Scandinavia show an ancestry similar to other contemporaneous farmer groups (Fig. 2A), with a majority of their ancestry related to early Neolithic farmers and a partial admixture component related to European Mesolithic HGs (Fig. 2B) (1, 2, 5⇓–7, 10, 16, 46). Fig. 2. (A) PCA of 429 present-day west Europeans (gray dots) with previously published Western HG (WHGs), Atlantic coast and Central European Neolithic farmer samples (filled symbols), and the samples from the present study (shaded symbols) projected onto the first two principal components (more details in SI Appendix, Section S11.1). (B) Inferred ancestry components (assuming seven clusters) of ancient individuals (Methods and SI Appendix, Section S11.2). All individuals to the left of Yoruba are prehistoric individuals, all of which are shotgun-sequenced unless marked with “CP” for SNP capture data. In the label names, the following letters indicate an archaeological context: CA, Chalcolithic; EN, Early Neolithic; N, Neolithic; MN, Middle Neolithic; LN, Late Neolithic. The LN individuals from Portugal come from different sites (key provided in Dataset S1.3). To further explore the demographic history of the individuals buried in the megaliths, we investigated the genetic affinities among sets of individuals and groups, using an f3 outgroup test for groups of individuals buried in megalithic or nonmegalithic contexts, as well as between individuals from Atlantic coastal and inland Neolithic sites (SI Appendix, section S11.3 and Fig. S19). These analyses showed genetic associations between individuals from the same/similar geographic region and time period (Fig. 2A and SI Appendix, Figs. S16 and S17). However, some tests (SI Appendix, Fig. S19) indicated similar trends as shown in our principal component analysis (PCA) and previous studies (5, 11, 15, 47, 48) and suggested a demic connection among western European Neolithic groups to the exclusion of central European Neolithic groups, as well as a connection between the British Isles and Iberian groups (SI Appendix, section S11.4 and Figs. S20–S22). These results were not driven by greater levels of HG ancestry among the populations at the fringes of the Neolithic expansion (11, 12, 15, 16) (SI Appendix, section S11.4). Interestingly, we also found a significant farmer-specific genetic affinity between the British Isles Neolithic populations and the Scandinavian populations (Ansarve and Gökhem; Fig. 1) to the exclusion of central European farmers (SI Appendix, Figs. S21 and S22). This observation is compatible with a further migration of farming groups along the European Atlantic coast, as has been suggested by the archaeological record (21, 49, 50). We found that significantly more males than females were buried in the British Isles megaliths (31 of 42 randomly sampled individuals; P = 0.0014, binomial test) and at the Primrose megalith alone (9 of 11; P = 0.032) (SI Appendix, section S8). However, other megalithic tombs with at least four individuals investigated, including Ansarve (6 of 9; P = 0.25), Gökhem (1 of 4; P = 0.93), La Mina (2 of 4; P = 0.68), Holm of Papa Westray (2 of 4; P = 0.68), and Isbister (Tomb of the Eagles) (8 of 10; P = 0.054), did not show the same striking pattern, nor did nonmegalithic burials from the British Isles (15) (nonmegalithic burials: 6 of 10; P = 0.27, cave burials: 10 of 15; P = 0.27, both nonmegalithic and cave burials: 16 of 25; P = 0.11). Overall, genetic data from all individuals from megalithic contexts suggest a higher male-to-female ratio in these burial chambers (41 of 60; P = 0.0031) (SI Appendix, Table S3), although the tendency is similar (but not significant) for nonmegalithic burials (SI Appendix, section S8). We found greater macrohaplogroup mtDNA diversity than Y-chromosomal (YDNA) diversity. Whereas mtDNA lineages from megalith burials harbor haplogroups K, H, HV, V, U5b, T, and J (among others), males from megalith burials belong almost exclusively to YDNA haplogroup I, more specifically to the I2a sublineage, which has a time to most recent common ancestor of ∼15000 BCE (51). This pattern of uniparental marker diversity is found not only among individuals buried in megaliths, but also in other farmer groups from the fourth millennium BCE, which display similar patterns of uniparental marker diversity (SI Appendix, Figs. S6 and S23) (10, 15, 48, 52). Some mtDNA lineages appear to be overrepresented at megalithic sites, with information from more than six individuals, including Primrose (n = 11; K1a+195 and K1a4a1 at 36% and 18% frequency, respectively), Ansarve (n = 9; J1c5 and K2b1a at ∼20% frequency), and Isbister (n = 10; K1a+195 at 20% frequency). Males from the present study belonged to YDNA haplogroup I, and those who could be resolved beyond this level were characterized as belonging to the I2a2a or I2a1b branch. Four of the 10 Primrose/Carrowmore males (Primrose 9, 12, 13, and 16) could be further resolved to the former sublineage, while the two Scottish males and the four Ansarve males could be further placed in the latter branch (Table 1 and SI Appendix, section S7). Combining the YDNA lineages and the radiocarbon dates of the individuals, a possible scenario of paternal continuity is observed for the Primrose and Ansarve megaliths. From the Primrose site, Primrose 9, 13, and 16, separated in time by at least 1 generation and possibly up to 12 generations, display the I2a2a1a1a haplotype. In addition, the Primrose 3, 10, and 17 individuals were inferred to harbor variants common to the I2a2 lineage, although with low coverage support (SI Appendix, section S7). A similar scenario is observed for the Ansarve megalith, with the individuals Ansarve 8, 14, and 17, separated by at most a few generations, carrying haplotype I2a1b1a. Ansarve 16, dated to at least 100 y younger, shares variants along the I2a1b lineage (Table 1 and SI Appendix, section S7).
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Post by Admin on Oct 16, 2019 18:23:47 GMT
The high frequency of the HG-derived I2a male lineages among megalith as well as nonmegalith individuals (SI Appendix, section S11.6) suggests a male sex-biased admixture process between the farmer and the HG groups (2, 12, 53, 54), but when this admixture occurred is unclear. To characterize the extent of sex-biased admixture between HGs and the individuals of the megalithic contexts, we assessed the affinity of all individuals buried in megaliths with sufficient genetic data, to an Early Neolithic farmer or a HG ancestry on the autosomes and the X chromosome using f4-statistics (SI Appendix, section S11.5). Higher levels of HG admixture on the autosomes than on the X chromosome implies a greater genetic contribution of male HGs than female HGs to these individuals, suggesting an HG male sex bias admixture. We find that in general, megalith groups do not harbor higher levels of HG ancestry on the autosomes compared with on the X chromosome (SI Appendix, Table S7 and Dataset S1.6), but the Scottish_MN farmers of this study showed a tendency toward an HG male-sex biased admixture in the recent past. The Scandinavian (Ansarve and Gökhem) individuals displayed an HG admixture for both the autosomes and the X chromosome (SI Appendix, Table S7), suggesting a scenario of more recent admixture with HGs in northern Europe. Using READ (Relationship Estimation from Ancient DNA) software (55), we inferred six kin relationships among the megalith individuals of this study: five relations among the Irish megaliths (two first-degree and three second-degree connections) and a second-degree relation in the Ansarve tomb (Fig. 3 and SI Appendix, section S10). First-degree relationships are characterized by either parent-offspring or a full sibling relationship, second-degree kin connections are represented by half-siblings, grandparent-grandchild, aunt/uncle-niece/nephew, and double cousins. Combining the READ predictions, uniparental lineages, radiocarbon dating, and age at death if available for those individuals who could be assessed, we inferred the potential familial relationships (Fig. 3 and SI Appendix, sections S2, S6, S7, and S10). Among the Irish megaliths, we observed two potential familial structures (SI Appendix, Fig. S10). The first is composed of three individuals from Primrose Grange (Tomb 1; individuals Primrose 2, 17, and 18), which overlap broadly in time (Fig. 3). Primrose 2 and 17 were predicted to be related in the first degree, representing a father-daughter relationship. Primrose 17 and 18 were predicted to be second-degree relatives (harboring the same mtDNA lineage but with possibly different YDNA haplogroups) and thus could have been half-siblings or double cousins. However, the YDNA prediction is hindered by low coverage and few informative markers, and thus a grandfather-grandson or uncle-nephew relationship cannot be fully excluded. Fig. 3. Kinship relationships in the Primrose, Carrowmore, and Ansarve burials. Solid line, first degree; dashed line, second degree. Males are displayed in green; females, in orange. The MtDNA and YDNA haplogroups are presented to the right of the figures. Bars underneath figures represent calibrated dating, with 95% CI (details in Table 1 and SI Appendix, Table S1). The other Irish putative pedigree structure was integrated by two individuals from Tomb 1 (Primrose 6 and 7) and one individual from Carrowmore 4 (from the Listhogil Tomb at the Carrowmore site in close vicinity), who harbored different mtDNA lineages. While the 95% CI dating range of Primrose 6 and Carrowmore 4 overlap, Primrose 7 might be slightly younger than the other two individuals. The Carrowmore 4 and Primrose 7 males were inferred to be at least second-degree related (3.14 SE below the expected value for two unrelated individuals), and the best prediction was a first-degree relation (1.79 SE below the value for a second-degree relation, although not statistically significant at the 95% level; SI Appendix, section S10). If a first-degree relation is assumed, then the sole possible kin connection is a father-son relationship, because the individuals are not maternally linked. In the case of a second-degree relationship, any paternally related second-degree familial connection is possible. The other two READ-predicted second-degree kin relationships in the Irish burials (Primrose 6-Primrose 7 and Primrose 6-Carrowmore 4; 1.04 SE and 0.50 SE below the threshold for an unrelated pair, respectively) involved a familial connection of the male individuals to Primrose 6 (female). Within the Ansarve megalith, we identify a second-degree relationship between the contemporaneous males Ansarve 14 and Ansarve 17 (Fig. 3 and SI Appendix, section S10). Both males have the same YDNA haplotype but different mtDNA lineages, suggesting that they could be related through any second-degree paternal kin relationship. Morphologically, Ansarve 14 was predicted to be an adult, and Ansarve 17 was predicted to be a juvenile (SI Appendix, section S2). Such observations might favor a grandfather-grandson or uncle-nephew relatedness over half-siblings or double cousins; however, the latter alternatives are still compatible with the data (SI Appendix, Fig. S12). READ analyses from other megalith burials where genetic data from at least four individuals were available per site (Gökhem, La Mina, Isbister, and Holm of Papa Westray; Fig. 1) did not reveal any evidence of genetic kinship relations. However, such observations may be hindered by the limited number of individuals investigated or by low genome coverage, which decreases the power to infer kinship (SI Appendix, section S10).
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Post by Admin on Oct 17, 2019 4:19:44 GMT
Discussion The genetic variation and characteristics of individuals buried in megalithic tombs, and also from individuals buried according to other traditions, suggest that the megalithic tradition was linked to socially stratified Neolithic farmer societies, with the genetic data suggesting close connections between Neolithic populations in Atlantic Europe (5, 15, 48) (Fig. 2 and SI Appendix, Figs. S19–S22). Here we provide evidence of a genetic connection among Scandinavian, British, and Irish Neolithic populations. This signal is weaker than the signals observed between the Iberian Peninsula and the British Isles, however (5, 11, 15, 47, 48) (Dataset S1.3), suggesting that migration between the British Isles and Scandinavia along the Atlantic coast was less frequent than that between Iberia and the British Isles (SI Appendix, section S11.4). The I2 YDNA lineages that are very common among European Mesolithic HGs (2, 3, 15, 56, 57) are distinctly different from the YDNA lineages of the European Early Neolithic farmer groups (8⇓–10), but frequent in the farmer groups of the fourth millennium BCE (2, 3, 8⇓–10, 15, 56, 57), suggesting a male HG admixture over time. The megalith individuals do not show higher levels of HG ancestry on the autosomes than on the X chromosome, but the Scottish_MN group shows a tendency toward a male-biased HG admixture in farmer groups, similar to previous observations (58). For the Scandinavian farmer groups, in contrast to the other megalith groups, we found an HG admixture for both the autosomes and the X chromosome. When these findings are considered together, it appears as if the social dynamics between HGs and Neolithic farmer groups, and thus the genetic admixture with HGs, differed somewhat in different geographic regions—an observation consistent with a combination of previous male sex bias admixture events occurring on the continent and more recent regional encounters with HG groups with a less pronounced sex-biased admixture. These observations imply that the groups that erected and used the megalithic burial structures were stable and stratified, but probably not isolated farmer societies (37, 41). The genetic connection of the individuals from the Primrose Grange and Carrowmore burials, spatially distanced by only 2 km and in contemporaneous use, suggests that transgenerational patrilineal structured societies could have expanded geographically, possibly leaving a (local) genetic fingerprint related to the social dynamics of the group. Such a scenario of forming patrilineal kin groups and intergroup competition during the Neolithic could explain the inferred Y-chromosome bottleneck seen in present-day European populations (51, 59). A central topic of discussion concerning the megalithic phenomena relates to the character of the communities that erected and used them for funerary rituals (27, 31, 37, 41, 42). The distinction of specific paternal lineages among the megaliths, a greater fraction of males than females in some megaliths, and their kindred relationships suggest that people buried in the megalithic tombs belonged to patrilineal segments of the groups/societies rather than representing a random sample from a larger Neolithic farmer community living in close vicinity. The sex ratio in the Irish megaliths is also in line with this finding. If one of the main functions of the tombs was to contain the remains of the deceased of a patrilineal segment, this would explain the inclusion of more males than females in the tombs. However, the finding that three of the five kinship relationships in these megaliths involved females indicates that female kindred members were not excluded. The observation of paternal continuity across time at the Gotlandic Ansarve megalith and at the Irish megaliths is a strong indication that specific family groups used these stone constructions for burial and other funerary practices. Of course, the patterns that we observe could be unique to the Primrose, Carrowmore, and Ansarve burials, and future studies of other megaliths are needed to provide additional data that can inform us further about social organization in the Neolithic. PNAS May 7, 2019 116 (19) 9469-9474
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Post by Admin on Nov 12, 2019 18:39:45 GMT
Fig. 1. Dolmen de las Ruines, Catalonia. Photo courtesy of B.S.P. There are ∼35,000 presently extant European megaliths, a term which is derived from Greek μέγας (mégas), “big,” and λίϑος (líthos), “stone.” These include megalithic tombs, standing stones, stone circles, alignments, and megalithic buildings or temples. Most of these were constructed during the Neolithic and the Copper Ages and are located in coastal areas. Their distribution is along the so-called Atlantic façade, including Sweden, Denmark, North Germany, The Netherlands, Belgium, Scotland, England, Wales, Ireland, northwest France, northern Spain, and Portugal, and in the Mediterranean region, including southern and southeastern Spain, southern France, the Islands of Corsica, Sardinia, Sicily, Malta and the Balearics, Apulia, northern Italy, and Switzerland. Interestingly, they share similar or even identical architectonic features throughout their distribution. Megalithic graves were built as dolmens and as passage or gallery graves (Figs. 1 and 2). Thousands of anthropogenic erected stones either stand isolated in the landscapes or were arranged as circles or in rows. There is evidence all across Europe for an orientation of the graves toward the east or southeast in the direction of the rising Sun. The question therefore arises whether there was a single, original source from which a megalithic movement spread over Europe or regional phenomena developed independently due to a similar set of conditions. Earlier research provided two very different answers to the question of origins. During the later 19th and the first two-thirds of the 20th centuries, archaeologists such as Montelius (1), Childe (2, 3), and Daniel (4) proposed models of a single origin of megaliths from which they then expanded by a process of diffusion. Thus, Montelius (1), in the Ex Oriente Lux Zeitgeist of the late 19th century, advocated for the Near East as a potential region of origin. Childe (5), building on Montelius, supported the idea of a diffusion of “oriental culture” by maritime exchange. According to Childe (6), the expansion was supported by a megalithic religion of migrant priestly elites who settled down long enough among local societies for the new ideas to take root. He proposed a route from the Mediterranean to the Atlantic northwest across the Pyrenean isthmus and an onward dissemination of the megalithic tradition from there to Britain and then later over the sea route around Spain and Portugal. Later, Childe (2, 3) expanded his theory about the spreading of a megalithic religion along the coastlines of western Europe by way of missionaries or prospectors. With the introduction of radiocarbon dates and processual approaches, the idea of an independent emergence of the same kind of stone architecture in several regions arose, because early C14 results did not support the diffusion model. Renfrew (7) was the first to exploit the new chronological results and proposed five independent nucleus centers, including Portugal, Andalusia, Brittany, southwest England, Denmark, and possibly Ireland for the emergence of megaliths in Europe. The model of an independent emergence of megaliths in several regions and sedentary, immobile farming communities has remained dominant in the research literature since then (8⇓–10). However, since the 1970s, the number of C14 dates of megaliths has expanded enormously. It is therefore timely to test the two prevailing interpretative models in the light of this new evidence. Fig. 2. Haväng dolmen, Scania. Strikingly, the architectonic concepts of megaliths are similar or even identical all over Europe. Photo courtesy of B.S.P. For this end, we investigated the fine-grain temporal pattern for the emergence of megaliths in Europe with the analysis of 2,410 available radiocarbon dates taken from premegalithic, megalithic, and nonmegalithic but contemporaneous contexts (Dataset S1). Radiocarbon dating is a two-stage process involving isotope measurements and the calibration against similar measurements made on dendrochronologically dated wood. For our time horizon, it normally provides precision ranges of 100 y to 300 y with 95% probability. To build a chronological megalithic sequence as precisely as possible, we adopted a Bayesian modeling approach, which is applied here to a wide region, using the program OxCal 4.1 (11, 12). We combined measurements with archaeological information relating to stratigraphical contexts, associated cultural material, and information on the burial rites, to narrow the time intervals for the calibrated ranges. In a first important step, we reviewed critically the 2,410 samples, including measurements from the 1960s up to the present, to determine the quality and reliability of the sample contexts. For each site with available radiocarbon results and a suitable sequence, we constructed one-phased or multiphased models with phase boundaries (Datasets S2 and S3) taking into consideration the detailed stratigraphic information (13). The posterior density estimates expressed as probability distributions in the text and in the figures are given by convention in italics to distinguish them clearly from simple calibrated radiocarbon dates. Results The radiocarbon dates suggest that the first megalithic graves in Europe were closed small structures or dolmens built aboveground with stone slabs and covered by a round or long mound of earth or stone. These graves emerge in the second half of the fifth millennium calibrated years (cal) BC within a time interval of 4794 cal BC to 3986 cal BC (95.4%; 4770 cal BC to 4005 cal BC, 68.2%) (Dataset S3, M7-2 to M29-4), which can be reduced most probably to 200 y to 300 y, in northwest France, the Channel Islands, Catalonia, southwestern France, Corsica, and Sardinia. Taking the associated cultural material into consideration, megalithic graves from Andalusia, Galicia, and northern Italy presumably belong to this first stage (Fig. 3). There are no radiocarbon dates available from the early megalithic graves in these regions, or their calibrated ranges show an onset extending into the fourth millennium cal BC, as is the case for Galicia. Of these regions, northwest France is the only one which exhibits monumental earthen constructions before the megaliths (SI Appendix, Fig. S2). The Passy graves in the Paris Basin have no megalithic chamber yet, but are impressive labor-intensive structures with a length of up to 280 m. These graves seem to be the earliest monumental graves in Europe; the first individual buried in the Passy necropolis died in 5061 cal BC to 4858 cal BC (95.4%; 5029 cal BC to 4946 cal BC, 68.2%) (Dataset S3, M1-4). Somewhat later, the first monumental graves emerge in Brittany, and especially in the region of Carnac, in the form of round tumuli covering pit burials, stone cists, and dry-wall chambers. The first building phase of the tumulus St. Michel in Carnac is dated to the time interval 4782 cal BC to 4594 cal BC (95.4; 4724 cal BC to 4618 cal BC, 68.2%) (Dataset S3, M4-2 to M4-4). The earliest megalithic grave chambers in Brittany, such as Tumiac, Kervinio, Castellic, St. Germain, Manio 5, Mané Hui, and Kerlescan (14⇓–16), emerge within this horizon as an architectonic feature of monumental long and round mounds. For these early megaliths, no radiocarbon determinations are available. It is only possible to limit the time interval of construction to the Ancient Castellic horizon based on the typochronological considerations of the grave goods and according to Ancient Castellic contexts with associated radiocarbon results ranging from 4794 cal BC to 3999 cal BC (95.4%; 4770 cal BC to 4034 cal BC, 68.2%) (Dataset S3, M7-2 to M7-7). Fig. 3. Map showing dates estimated for the start of megaliths in the different European regions, with 95% probability (68% probability in brackets). Italic bold type is used for date ranges of the posterior density estimates based on samples from megalithic contexts, regular bold type is used for simple calibrated radiocarbon dates from megalithic contexts, and regular italic type is used for the probabilities of the posterior density estimates associated with the earliest cultural material in the megaliths. In Catalonia, in the Tavertet region, early megalithic graves emerged during the same time interval, even contemporaneous with the graves in Brittany. A reevaluation of the available radiocarbon results yielded a dating of the construction of these graves not before 4722 cal BC to 4068 cal BC (95.4%; 4581 cal BC to 4267 cal BC, 68.2%) (Dataset S3, M24-33). A part of these data exhibit an inbuilt age (Dataset S3, M24-28 to M24-32) (ref. 13, p. 128). On the northeastern side of the Pyrenees in southern France, early megaliths are either isolated in the landscape or arranged in necropolises as at Najac and Camp del Ginèbre. The unmodeled ranges of three radiocarbon results for human bones from the necropolis of Najac 4328 cal BC to 3979 cal BC (95.4%; 4318 cal BC to 3995 cal BC) (Dataset S1, 830 to 832) suggest burials within this time horizon. Along the central Mediterranean coasts and north Mediterranean islands of Sardinia and Corsica, small necropolises are found with early megalithic graves. The grave goods from the Li Muri necropolis on Sardinia are attributed to the Late Neolithic San Ciriaco horizon, and, according to the radiocarbon results from the San Ciriaco layers in the settlement of Contraguda, it is possible to limit the emergence of these graves to a time interval from 4733 cal BC to 3986 cal BC (95.4%; 4471 cal BC to 4005 cal BC, 68.2%) (Dataset S3, M29-1 to M29-4). There are further clusters with potential early megalithic graves documented in the central Mediterranean in northern Italy, for example, in La Vela-Trento, or Maddalena di Chiomonte-Torino and possibly Apulia (6). However, for these, there are no radiocarbon dates available yet. Based on the archaeological material, they are likely dated to the second half of the fifth millennium cal BC. From the southwest Iberian Peninsula in Andalusia, the Algarve, and the Alentejo, we find more of these possible early megaliths (17⇓–19). In the northern half of the western Iberian Peninsula, there are early megaliths, concentrated mainly in Galicia. So far, these have been dated to the very end of the fifth millennium cal BC, if not later. Most of these dates are from charcoal, and many represent termini post quos values due to the inbuilt age of the wood or unsure contexts. From Chan de Cruz 1, a possible construction or usage date from ∼4080 cal BC (CSIC-642, 5210 ± 50 BP, 4144 cal BC to 3961 cal BC, 68.2%; 4230 cal BC to 3947 cal BC, 95.4%) (Dataset S1, 2014) is available.
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