Uralic speakers formed clines of Corded Ware ancestry with WHG:ANE populations

steppe-forest-tundra-biomes-uralic

The preprint by Jeong et al. (2018) has been published: The genetic history of admixture across inner Eurasia Nature Ecol. Evol. (2019).

Interesting excerpts, referring mainly to Uralic peoples (emphasis mine):

A model-based clustering analysis using ADMIXTURE shows a similar pattern (Fig. 2b and Supplementary Fig. 3). Overall, the proportions of ancestry components associated with Eastern or Western Eurasians are well correlated with longitude in inner Eurasians (Fig. 3). Notable outliers include known historical migrants such as Kalmyks, Nogais and Dungans. The Uralic- and Yeniseian-speaking populations, as well as Russians from multiple locations, derive most of their Eastern Eurasian ancestry from a component most enriched in Nganasans, while Turkic/Mongolic speakers have this component together with another component most enriched in populations from the Russian Far East, such as Ulchi and Nivkh (Supplementary Fig. 3). Turkic/Mongolic speakers comprising the bottom-most cline have a distinct Western Eurasian ancestry profile: they have a high proportion of a component most enriched in Mesolithic Caucasus hunter-gatherers and Neolithic Iranians and frequently harbour another component enriched in present-day South Asians (Supplementary Fig. 4). Based on the PCA and ADMIXTURE results, we heuristically assigned inner Eurasians to three clines: the ‘forest-tundra’ cline includes Russians and all Uralic and Yeniseian speakers; the ‘steppe-forest’ cline includes Turkic- and Mongolic-speaking populations from the Volga and Altai–Sayan regions and Southern Siberia; and the ‘southern steppe’ cline includes the rest of the populations.

eurasian-clines-uralic-altaic
The first two PCs summarizing the genetic structure within 2,077 Eurasian individuals. The two PCs generally mirror geography. PC1 separates western and eastern Eurasian populations, with many inner Eurasians in the middle. PC2 separates eastern Eurasians along the northsouth cline and also separates Europeans from West Asians. Ancient individuals (color-filled shapes), including two Botai individuals, are projected onto PCs calculated from present-day individuals.

For the forest-tundra populations, the Nganasan + Srubnaya model is adequate only for the two Volga region populations, Udmurts and Besermyans (Fig. 5 and Supplementary Table 8).

For the other populations west of the Urals, six from the northeastern corner of Europe are modelled with additional Mesolithic Western European hunter-gatherer (WHG) contribution (8.2–11.4%; Supplementary Table 8), while the rest need both WHG and early Neolithic European farmers (LBK_EN; Supplementary Table 2). Nganasan-related ancestry substantially contributes to their gene pools and cannot be removed from the model without a significant decrease in the model fit (4.1–29.0% contribution; χ2 P ≤ 1.68 × 10−5; Supplementary Table 8).

west-urals-finno-ugrians-qpadm
Supplementary Table 8. QpAdm-based admixture modeling of the forest-tundra cline populations. For the 13 populations west of the Urals, we present a four-way admixture model, Nganasan+Srubnaya+WHG+LBK_EN, or its minimal adequate subset. Modified from the article, to include colors for cultures, and underlined best models for Corded Ware ancestry among Uralians.

NOTE. It doesn’t seem like Hungarians can be easily modelled with Nganasan ancestry, though…

For the 4 populations east of the Urals (Enets, Selkups, Kets and Mansi), for which the above models are not adequate, Nganasan + Srubnaya + AG3 provides a good fit (χ2 P ≥ 0.018; Fig. 5 and Supplementary Table 8). Using early Bronze Age populations from the Baikal Lake region (‘Baikal_EBA’; Supplementary Table 2) as a reference instead of Nganasan, the two-way model of Baikal_EBA + Srubnaya provides a reasonable fit (χ2 P ≥ 0.016; Supplementary Table 8) and the three-way model of Baikal_EBA + Srubnaya + AG3 is adequate but with negative AG3 contribution for Enets and Mansi (χ2 P ≥ 0.460; Supplementary Table 8).

east-urals-ugric-samoyedic-qpadm
Supplementary Table 8. QpAdm-based admixture modeling of the forest-tundra cline populations. For the four populations east of the Urals, we present three admixture models: Baikal_EBA+Srubnaya, Baikal_EBA+Srubnaya+AG3 and Nganasan+Srubnaya+AG3. For each model, we present qpAdm p-value, admixture coefficient estimates and associated 5 cM jackknife standard errors (estimate ± SE). Modified from the article, to include colors for cultures, and underlined best models for Corded Ware ancestry among Uralians.

Bronze/Iron Age populations from Southern Siberia also show a similar ancestry composition with high ANE affinity (Supplementary Table 9). The additional ANE contribution beyond the Nganasan + Srubnaya model suggests a legacy from ANE-ancestry-rich clines before the Late Bronze Age.

bronze-age-iron-age-karasuk-mezhovska-tagar-qpadm
Supplementary Table 9. QpAdm-based admixture modeling of Bronze and Iron Age populations of southern Siberia. For ancieint individuals associated with Karasuk and Tagar cultures, Nganasan+Srubnaya model is insufficient. For all five groups, adding AG3 as the third ancestry or substituting Nganasan with Baikal_EBA with higher ANE affinity provides an adequate model. For each model, we present qpAdm p-value, admixture coefficient estimates and associated 5 cM jackknife standard errors (estimate ± SE). Models with p-value ≥ 0.05 are highlighted in bold face. Modified from the article, to include colors for cultures, and underlined best models for Corded Ware ancestry among Uralians.

Lara M. Cassidy comments the results of the study in A steppe in the right direction (you can read it here):

Even among the earliest available inner Eurasian genomes, east–west connectivity is evident. These, too, form a longitudinal cline, characterized by the easterly increase of a distinct ancestry, labelled Ancient North Eurasian (ANE), lowest in western European hunter-gatherers (WHG) and highest in Palaeolithic Siberians from the Baikal region. Flow-through from this ANE cline is seen in steppe populations until at least the Bronze Age, including the world’s earliest known horse herders — the Botai. However, this is eroded over time by migration from west and east, following agricultural adoption on the continental peripheries (Fig. 1b,c).

Strikingly, Jeong et al. model the modern upper steppe cline as a simple two-way mixture between western Late Bronze Age herders and Northeast Asians (Fig. 1c), with no detectable residue from the older ANE cline. They propose modern steppe peoples were established mainly through migrations post-dating the Bronze Age, a sequence for which has been recently outlined using ancient genomes. In contrast, they confirm a substantial ANE legacy in modern Siberians of the northernmost cline, a pattern mirrored in excesses of WHG ancestry west of the Urals (Fig. 1b). This marks the inhospitable biome as a reservoir for older lineages, an indication that longstanding barriers to latitudinal movement may indeed be at work, reducing the penetrance of gene flows further south along the steppe.

eurasian-clines-uralic-turkic-mongol-altaic
The genomic formation of inner Eurasians. b–d, Depiction of the three main clines of ancestry identified among Inner Eurasians. Sources of admixture for each cline are represented using proxy ancient populations, both sampled and hypothesised, based on the study’s modelling results. The major eastern and western ancestries used to model each cline are shown in bold; the peripheral admixtures that gave rise to these are also shown. Additional contributions to subsections of each cline are marked with dashed lines. b, The northernmost cline, illustrating the legacy of WHG and ANE-related populations. c,d, The upper (c) and lower (d) steppe clines are shown, both of which have substantial eastern contributions related to modern Tungusic speakers. The authors propose these populations are themselves the result of an admixture between groups related to the Nganasan, whose ancestors potentially occupied a wider range, and hunter-gatherers (HGs) from the Amur River Basin. While the upper steppe cline in c can be described as a mixture between this eastern ancestry and western steppe herders, the current model for the southern steppe cline as shown in d is not adequate and is likely confounded by interactions with diverse bordering ancestries. Credit: Ecoregions 2017, Resolve https://ecoregions2017.appspot.com/

Given the findings as reported in the paper, I think it should be much easier to describe different subclines in the “northernmost cline” than in the much more recent “Turkic/Mongolic cline”, which is nevertheless subdivided in this paper in two clines. As an example, there are at least two obvious clines with “Nganasan-related meta-populations” among Uralians, which converge in a common Steppe MLBA (i.e. Corded Ware) ancestry – one with Palaeo-Laplandic peoples, and another one with different Palaeo-Siberian populations:

siberian-clines-uralic-altaic
PCA of ancient and modern Eurasian samples. Ancient Palaeo-Laplandic, Palaeosiberian, and Altai clines drawn, with modern populations labelled. See a version with higher resolution.

The inclusion of certain Eurasian groups (or lack thereof) in the PCA doesn’t help to distinguish these subclines visually, and I guess the tiny “Naganasan-related” ancestral components found in some western populations (e.g. the famous ~5% among Estonians) probably don’t lend themselves easily to further subdivisions. Notice, nevertheless, the different components of the Eastern Eurasian source populations among Finno-Ugrians:

uralic-admixture-qpadm
Characterization of the Western and Eastern Eurasian source ancestries in inner Eurasian populations. [Modified from the paper, includes only Uralic populations]. a, Admixture f3 values are compared for different Eastern Eurasian (Mixe, Nganasan and Ulchi; green) and Western Eurasian references (Srubnaya and Chalcolithic Iranians (Iran_ChL); red). For each target group, darker shades mark more negative f3 values. b, Weights of donor populations in two sources characterizing the main admixture signal (date 1 and PC1) in the GLOBETROTTER analysis. We merged 167 donor populations into 12 groups (top right). Target populations were split into five groups (from top to bottom): Aleuts; the forest-tundra cline populations; the steppe-forest cline populations; the southern steppe cline populations; and ‘others’.

Also remarkable is the lack of comparison of Uralic populations with other neighbouring ones, since the described Uralic-like ancestry of Russians was already known, and is most likely due to the recent acculturation of Uralic-speaking peoples in the cradle of Russians, right before their eastward expansions.

west-eurasian-east-eurasian-ancestry
Supplementary Fig. 4. ADMIXTURE results qualitatively support PCA-based grouping of inner Eurasians into three clines. (A) Most southern steppe cline populations derive a higher proportion of their total Western Eurasian ancestry from a source related to Caucasus, Iran and South Asian populations. (B) Turkic- and Mongolic-speaking populations tend to derive their Eastern Eurasian ancestry more from the Devil’s Gate related one than from Nganasan-related one, while the opposite is true for Uralic- and Yeiseian-speakers. To estimate overall western Eurasian ancestry proportion, we sum up four components in our ADMIXTURE results (K=14), which are the dominant components in Neolithic Anatolians (“Anatolia_N”), Mesolithic western European hunter-gatherers (“WHG”), early Holocene Caucasus hunter-gatherers (“CHG”) and Mala from southern India, respectively. The “West / South Asian ancestry” is a fraction of it, calculated by summing up the last two components. To estimate overall Eastern Eurasian ancestry proportion, we sum up six components, most prevalent in Surui, Chipewyan, Itelmen, Nganasan, Atayal and early Neolithic Russian Far East individuals (“Devil’s Gate”). Eurasians into three clines. (A) Most southern steppe cline populations derive a higher proportion of their total Western Eurasian ancestry from a source related to Caucasus, Iran and South Asian populations. (B) Turkic- and Mongolic-speaking populations tend to derive their Eastern Eurasian ancestry more from the Devil’s Gate related one than from Nganasan-related one, while the opposite is true for Uralic- and Yeiseian-speakers. To estimate overall western Eurasian ancestry proportion, we sum up four components in our ADMIXTURE results (K=14), which are the dominant components in Neolithic Anatolians (“Anatolia_N”), Mesolithic western European hunter-gatherers (“WHG”), early Holocene Caucasus hunter-gatherers (“CHG”) and Mala from southern India, respectively. The “West / South Asian ancestry” is a fraction of it, calculated by summing up the last two components. To estimate overall Eastern Eurasian ancestry proportion, we sum up six components, most prevalent in Surui, Chipewyan, Itelmen, Nganasan, Atayal and early Neolithic Russian Far East individuals (“Devil’s Gate”).

A comparison of Estonians and Finns with Balts, Scandinavians, and Eastern Europeans would have been more informative for the division of the different so-called “Nganasan-like meta-populations”, and to ascertain which one of these ancestral peoples along the ancient WHG:ANE cline could actually be connected (if at all) to the Cis-Urals.

Because, after all, based on linguistics and archaeology, geneticists are not supposed to be looking for populations from the North Asian Arctic region, for “Siberian ancestry”, or for haplogroup N1c – despite previous works by their peers – , but for the Bronze Age Volga-Kama region…

Related

The origin of social complexity in the development of the Sintashta culture

kamenni-ambar

Very interesting PhD thesis by Igor Chechushkov, Bronze Age human communities in the Southern Urals steppe: Sintashta-Petrovka social and subsistence organization (2018).

Abstract:

Why and how exactly social complexity develops through time from small-scale groups to the level of large and complex institutions is an essential social science question. Through studying the Late Bronze Age Sintashta-Petrovka chiefdoms of the southern Urals (cal. 2050–1750 BC), this research aims to contribute to an understanding of variation in the organization of local communities in chiefdoms. It set out to document a segment of the Sintashta-Petrovka population not previously recognized in the archaeological record and learn about how this segment of the population related to the rest of the society. The Sintashta-Petrovka development provides a comparative case study of a pastoral society divided into sedentary and mobile segments.

Subsurface testing on the peripheries of three Sintashta-Petrovka communities suggests that a group of mobile herders lived outside the walls of the nucleated villages on a seasonal basis. During the summer, this group moved away from the village to pasture livestock farther off in the valley, and during the winter returned to shelter adjacent to the settlement. This finding illuminates the functioning of the year-round settlements as centers of production during the summer so as to provide for herd maintenance and breeding and winter shelter against harsh environmental conditions.

The question of why individuals chose in this context to form mutually dependent relationships with other families and thus give up some of their independence can be answered with a combination of two necessities: to remain a community in a newly settled ecological niche and to protect animals from environmental risk and theft. Those who were skillful at managing communal construction of walled villages and protecting people from military threats became the most prominent members of the society. These people formed the core of the chiefdoms but were not able to accumulate much wealth and other possessions. Instead, they acquired high social prestige that could even be transferred to their children. However, this set of relationships did not last longer than 300 years. Once occupation of the region was well established the need for functions served by elites disappeared, and centralized chiefly communities disintegrated into smaller unfortified villages.

sintashta-petrovka-archaeological
Research area: map of the Sintashta-Petrovka archaeological sites. Settlements: 101 – Stepnoye; 102 – Shibaeyvo 1; 103 – Chernorechye 3; 104 – Bakhta; 105 – Paris; 106 – Isiney; 107 – Kuisak; 108 – Ust’ye; 109 – Rodniki; 110 – Konoplyanka; 111 – Zhurumbay; 112 – Arkaim; 113 – Sintashta; 114 – Sintashta 2; 115 – Kamennyi Ambar; 116 – Alandskoye; 117 – Chekatay; 118 – Selek; 119 – Sarym- Sakly; 120 – Kamysty; 121 – Kizilskoye; 122 – Bersuat; 123 – Andreyevskoe; 124 – Ulak; 125 – Streletskoye; 126 – Zarechnoye 4; 127 – Kamennyi Brod. Cemeteries: 201 – Ozernoye 1; 202 – Krivoe Ozero; 203 – Stepnoye M; 204 – Kamennyi Ambar-5; 205 – Stepnoye 1; 206 – Tsarev Kurgan; 207 – Ubagan 2; 208 – Solntse 2; 209 – Bolshekaraganskyi; 210 – Aleksandrovsky 4; 211 – Sintashta; 212 – Solonchanka 1a; 213 – Knyazhenskyi; 214 – Bestamak; 215 – Ishkinovka 1; 216 – Ishkinovka 2; 217 – Novo–Kumakskyi; 218 – Zhaman–Kargala 1; 219 – Tanabergen 2; 220 – Novo-Petrovka; 221 – Semiozernoye 2; 222 – Khalvayi 3

Some interesting excerpts (emphasis mine):

The quintessential archaeological evidence of Sintashta-Petrovka communities takes the form of highly nucleated and fortified settlements paired with easily-recognized kurgan (burial mound) cemeteries. This pattern spread across Northern Central Eurasia in a relatively short period of about 300 years (cal. 2050–1750 BC), and the period consists of two chronological phases (Hanks et al. 2007). The earlier Sintashta phase (cal. 2050–1850 BC) is distinguished from the later Petrovka phase (cal. 1850–1750 BC) by some differences in ceramic styles and some techniques of bronze metallurgy (Degtyareva et al. 2001; Vinogradov 2013). Bronze Age subsistence patterns apparently relied on a wide variety of resources, among which meat and milk production played a major role (…). The most outstanding graves are individual male burials accompanied by weaponry (projectile weapons and chariots), the insignia of power (stone mace heads), craft tools, and a specific set of sacrificed animals (horses, cows, and dogs). (…) there were at least two adults buried with chariots and one with sacrificed horses (Epimakhov 1996b). Chariots – the most famous and spectacular material component of Sintashta-Petrovka society – are known exclusively from burial contexts. Two-wheeled vehicles represent complex technology, incorporating some crucial innovations and the investment of substantial resources. Highly developed craft and military skills were required for their production and use. Burials with chariots probably represent military elites who used them (Anthony 2009; Chechushkov 2011; Frachetti 2012:17) and played especially important social roles in Sintashta-Petrovka societies. This pattern strongly suggests that military leadership extended into the realm of ideology and general social prestige (Earle 2011:32–33).

The following sequence of archaeological cultures – based on the sample of radiocarbon dates (Epimakhov 2007a; 2010a), – is adopted: (1) the Sintashta-Petrovka phase 1 dated to cal. 2050–1750 BC and (2) the Srubnaya-Alakul’ phase 2 dated to cal. 1750–1350 BC.

(…) control of craft might have provided a source of power for elites in the fortified settlements (Steponaitis 1991). Some bronze tools, such as chisels, adzes, and handsaws seem more abundantly represented at some fortified settlements than at others, raising the possibility of a stronger focus on different craft products and some degree of exchange and interdependence between fortified settlements. (…) Zdanovich (1995:35) estimates 2500 people within the walls at Arkaim. He bases his conclusion an average house size of 140 m2 and the idea that Arkaim households consisted of an extended family of several generations, similar to Iroquois longhouse inhabitants. He also suggests that the entire population did not live in the “town” all the time, but moved around. The fully permanent residents were shamans, warriors, and craftsmen, i.e., elites and attached specialists.

Summarizing, excavated households represent very strongly similar architectural patterns, similar levels of wealth and prestige, little productive differentiation, and no evidence of elites amassing wealth through control of craft or subsistence production or any other mechanism (Earle 1987). These observations sharply contradict the burial record, where strong social differentiation is visible. The description above recalls the Regional Classic period elites of the Alto Magdalena whose standard of living differed little if at all from anyone else’s. Their elaborate tombs and sculptures suggest supernatural powers and ritual roles were much more important bases of their social prominence than economic control or accumulation of wealth (Drennan 1995:96–97). On the other hand, craft activities (especially metal production) are highly obvious in the Sintashta-Petrovka settlements. Defensive functions could also have played some role for the entire population. This benefit might attract people in an unstable or wild environment to spend much of their time in or near such settlements (Earle 2011:32–33). Since the construction of ditches and outer walls, as well as dwellings with shared walls, requires planning and organization, purposeful collective effort must have been a key feature of Sintashta-Petrovka communities (Vinogradov 2013; Zdanovich 1995). Sintashta-Petrovka communities thus evidence substantial investment of effort in non-subsistence activities, potentially resulting in a subsistence deficit in an economy with a heavy emphasis on herding. Altogether, this makes it plausible to think of the known Sintashta-Petrovka communities as special places where elites for whom military activities were important resided, and where metal production and possibly other crafts were carried out. It remains unclear just how a subsistence economy relying heavily on herding was managed from these substantial sedentary communities. Moving herds around the landscape seasonally is generally thought to be a part of subsistence strategy in Inner Eurasia (Frachetti 2008; Bachura 2013). In this area migration to exploit seasonal pastures is the best strategy for maintaining a regular supply of food for livestock due to shortages of capital or of labor pool to produce, harvest, and store fodder (Dyson-Hudson and Dyson-Hudson 1980:17). The recent stable isotope studies support this notion showing high likelihood that during the Bronze Age livestock was raised locally (Kiseleva et al. 2017).

The above raises the possibility that the residential remains that have been excavated within the fortifications of Sintashta-Petrovka communities represent only a portion of the population (Hanks and Doonan 2009, Johnson and Hanks 2012). It could be (along with the general lines suggested by D. Zdanovich [1997]) that the archaeological remains of the ordinary people who made up the majority of the population, built the impressive fortifications and stoked the subsistence economy have gone largely undetected. In global comparative perspective, many societies with the features known for Sintashta-Petrovka organization consisted of elite central-place settlements and hinterland populations. In such a scenario, the “missing” portion of the Sintashta population would reside in smaller unfortified settlements scattered around in the vicinity of the fortified ones.

kamenni-ambar-cultural-layer

In terms of wealth and productive differentiation, the inside assemblage of Kamennyi Ambar demonstrates a higher degree of richness and diversity in its material assemblage, leading to the conclusion that the outside materials may represent a semi-mobile group of people who used significantly less durable materials and accumulated less possessions. As for the diversity within the inside artifact assemblage, some households at Kamennyi Ambar demonstrate more diverse artifact assemblages than others, as well as bigger sizes, that could be related to differences in productive activities and/or wealth differentiation between families. A focus on specific objects of ceramic production in House 1 suggests some degree of productive specialization, while the elite goods in House 5 clearly point out the presence of elite members of the society.

There are two possible social scenarios that explain the settlement situation during the Sintashta-Petrovka phase. The first scenario considers all three communities as simultaneous and the second scenario suggests seeing the three sites as the same community that moved around the landscape during the Late Bronze Age in order to keep the pasture grounds from degradation.

Since no remains of permanent structures were found and any people living outside the walls must have stayed in temporary shelters. If this was the case, then the outside part of the population consisted of a semi-mobile group of people who moved to live near the fortified settlement during the winter. The pattern of animal slaughtering supports this conclusion. Animal teeth found near Kamennyi Ambar and Konoplyanka demonstrate a tendency for animal butchering during the fall, throughout the winter and spring, with less evidence of summer meat consumption. Moreover, since the Bronze Age subsistence strategy relied heavily on pastoralism, herds had to be grazed during the summer and kept safe during the winter. This strongly suggests that the part of the population responsible for management of animals spent their time in the summer pastures with the livestock. During the winter the animals had to be kept in the warm and safe environment of the walled settlements (as suggested by the highest level of phosphorus on the house floors) while the herders stayed in portable shelters in close to the walls.

(…) the outsiders used a less diverse set of tools, as well as less durable materials (for example, wooden instead of metal) in their everyday life and did not accumulate much in the way of archaeologically visible possessions. On the other hand, a few stone and lithic artifacts demonstrate that craft activities were carried out using cheap and abundant raw materials. The artefact assemblages also point out that the people inside accumulated wealth in the form of material belongings and luxury goods, especially, things like metal artifacts and symbolic or military-related stone artifacts, while people outside did not do that. However, the presence of semi-precious stones could signify some kind of wealth accumulation by the segment of population outside the walls. Since there are limits to our ability to assess social relationships from material remains, it is difficult to say if the people who lived outside the walls were oppressed or less respected. Their possible concentration on herding-related activities and livestock keeping might suggest less prestigious social status. The most prominent members of the society were, nonetheless, buried with the attributes of warriors or craft specialists, not those of shepherds, suggesting that those involved in livestock management had less social prestige.

Furthermore, Kuzmina (1994:72) cites linguistic studies demonstrating that the Sanskrit word for a permanent village earlier meant a circle of mobile wagon homes, situated together for defensive purposes for an overnight camp (Kuzmina 1994:72).

The likely population of semi-mobile herders represented some 30%–60% of the entire local community, while the other of 40%–70% were inhabitants of the walled settlement. The almost completely excavated kurgan cemetery of Kamennyi Ambar-5 (only two kurgans remain unstudied) yielded about 100 individuals, or about 2%–5% of the total of 4,896±1,960 individuals in four generations who lived at the nearby settlement for 100 years. In other words, no more than 10% of the population was entitled to be buried under the kurgan mound and this proportion can be taken as an estimate of those with elevated social status. Perhaps, these elites were kin, since analysis of the burial patterns suggests sex/age rather than wealth/prestige differentiation between buried individuals within this elite group (Epimakhov and Berseneva 2011; Ventresca Miller 2013). The remaining non-elite members of the permanently resident community, then, represented some 30%–60% of the complete local community, but did not show evidence of standards of living particularly lower than the elites eventually interred in the kurgan.

(…) The buried population in the Sintashta Cemetery is about 80 individuals or only about 2%–3% of the total estimated population. However, these few individuals were buried with extremely rich offerings, like complete chariots, decorations made of precious metals or sacrifices of six horses (equal to about 900 kg of meat), etc. With such a low proportion of the population assigned such high prestige, the Sintashta local community can easily be labeled a local chiefdom. In Pitman and Doonan’s view (2018) the social structure of the chifedom consisted of a chief and his kin at the highest level; warriors, religious specialists, and craftsmen in the middle; and the pastoral community at the bottom level.

kamenni-ambar-excavations

In the Bronze Age, the people who comprised the majority of the permanent population were involved in craft activities, including extraction of copper ores, metallurgy, bone, leather, and woodwork. The most important and labor-intensive part of the economy, however, was haymaking. The evidence of hay found in the cultural layer near Kamennyi Ambar supports the idea that animals were fed during the winter. Nowadays, hay cutting is typically done in July-August, the period of most intensive grazing for animals. Thus, the part of the collective that remained in the settlement had to provide the labor force for haymaking.

In the wintertime, the herders returned to the settlements with the herds, and animals were kept inside the walls––a practice which is known archaeologically (Zakh 1995) and ethnographically (Shahack-Gross et al. 2004)––while herders stayed outside in their tents.

In sum, the Sintashta-Petrovka chiefdoms demonstrate a three-part social order. In Kuzmina’s (1994) view, this is similar to the Varna system of ancient India, that consisted of priests (Sansk. Brahmanis), rulers and warriors (Sansk. Kshatriyas), free producers (Sansk. Vaishyas) and laborers and service providers (Sansk. Shudras). In the Sintashta-Petrovka chiefdom, the elite 2%–5% of the population would have consisted of priests and warriors; 48%–55% would have been dependent producers; and 50%–60% would have been herders of lower social rank.

sintashta-petrovka-settlements
The map of the Bronze Age sites in the Karagaily-Ayat Valley Sites of Phase 1: 101 – Konoplyanka; 102 – Zhurumbay; 103 – Kamennyi Ambar; 104 – Kamennyi Ambar-5 Sites of Phase 2: 201 – Konoplyanka 1; 202 – Varshavskoye-1; 203 – Zhurumbay-1; 204 – Varshavskoye-3; 205 – Varshavskoye-5; 206 – Varshavskoye-9; 207 – Kamennyi Ambar-8; 208 – Kamennyi Ambar; 209 – Elizavetpolskoye-3; 210 – Elizavetpolskoye-2; 211 – Karagayli-26; 212 – Elizavetpolskoye-7; 213 – Elizavetpolskoye- 9; 214 – Yuzhno-Stepnoyi (1); 215 – Yuzhno-Stepnoyi (2)

Conclusions

In the case of the Sintashta-Petrovka chiefdoms, the questions of why and how exactly social complexity developed through time and why individuals choose to integrate and give up their independence can be answered as some combination of two necessities: to persist as a larger community in the ecological niche of the newly settled region, and to protect herds from theft.

There is general agreement among researchers that the Sintashta phenomenon had no local roots and originated with a large-scale migration of pastoral communities from Eastern Europe to the marginal area of the Southern Urals. This process forced families to stay together and fueled the necessity in the walled villages for ensuring the reproduction of herds in the extreme climatic conditions of the southern Urals that are colder and dryer than the eastern Black Sea region from which the Sintashta populations are thought to have migrated (Kuzmina 1994, 2007; Anthony 2007; Vinogradov 2011, etc.). At the same time, the herds needed protection from animal and human predators. Probably, the risk of losing animals was a threat to survival that created tensions between neighboring communities, and the Neolithic hunter-gatherers who had populated the Urals before the arrival of Sintashta people could have hunted the domestic animals. Apparently, those who were talented in managing the construction of closely-packed villages surrounded by ditches and walls to protect people and livestock from threats from neighbors, and who otherwise served the community in the newly colonized zone became the most prominent members of society. Theses people formed the core of the Sintashta-Petrovka chiefdom but were not able to accumulate much personal wealth in the form of material possessions. Instead, they acquired high social prestige that could even be transferred to their children (since up to 65% of the buried elite population consists of infants [Razhev and Epimakhov 2005). In this sense, the Sintashta-Petrovka elites were simmilar to their counterparts in the Alto Magdalena of Colombia (Drennan 1995; Gonzalez Fernandez 2007; Drennan and Peterson 2008).

However, this situation did not last longer than 300 years, since after the initial phase of colonization of the Southern Urals was over, the need for social services provided by an elite disappeared and centralized chiefly communities disintegrated into the smaller unfortified villages of the Srubnaya-Alakul’ period.

As I have said many times already (see e.g. here) the outsider pastoralists, forming originally the vast majority of the population, were most likely Pre-Proto-Indo-Iranian speakers of haplogroup R1b-Z2103, and their elite groups (whose inheritance system was based on kinship) probably incorporated gradually Uralic-speaking families of haplogroup R1a-Z93, whose relative importance increased gradually, and then eventually expanded massively with the migrations of Andronovo and Srubna, creating a second Y-chromosome bottleneck that favoured again Z93 subclades. The adaptation of Pre-Proto-Indo-Iranian to the Uralic pronunciation, and the adoption of PII vocabulary in neighbouring Proto-Finno-Ugric bear witness to this process.

Related

Sintashta diet and economy based on domesticated animal products and wild resources

indo-iranian-sintashta-uralic-migrations

New paper (behind paywall) Bronze Age diet and economy: New stable isotope data from the Central Eurasian steppes (2100-1700 BC), by Hanks et al. J. Arch. Sci (2018) 97:14-25.

Interesting excerpts (emphasis mine):

Previous research at KA-5 was carried out by A. V. Epimakhov in 1994–1995 and 2002–2003 and resulted in the excavation of three Sintashta culture barrows (kurgans) that produced 35 burial pits and a reported 100 skeletons (Epimakhov, 2002, 2005; Epimakhov et al., 2005; Razhev and Epimakhov, 2004). Seven AMS radiocarbon dates on human remains from the cemetery yielded a date range of 2040–1730 cal. BC (2 sigma), which placed the cemetery within the Sintashta phase of the regional Bronze Age (Hanks et al., 2007). Twelve recently obtained AMS radiocarbon dates, taken from short-lived wood and charcoal species recovered from the Kamennyi Ambar settlement, have provided a date range of 2050–1760 cal. BC (2 sigma). Importantly, these dates confirm the close chronological relationship between the settlement and cemetery for the Middle Bronze Age phase and discount the possibility of a freshwater reservoir effect influencing the earlier dating of the human remains from the Kamennyi Ambar 5 cemetery (Epimakhov and Krause, 2013).

Sintashta cemeteries frequently yield fewer than six barrow complexes and the number of skeletons recovered represents a fraction of the total population that would have inhabited the settlements (Judd et al., 2018; Johnson and Hanks, 2012). Scholars have suggested that only members of higher status were afforded interment in these cemeteries and that principles of social organization structured placement of individuals within central or peripheral grave pits (Fig. 2) (Koryakova and Epimakhov, 2007: 75–81). In comparison with other Sintashta cemeteries that have been excavated, KA-5 provides one of the largest skeletal inventories currently available for study.

kamenniy-ambar
Upper – plan of Kamennyi Ambar settlement and cemetery; Lower – plan views of Kurgan 2 and Kurgan 4 from KA-5 Cemetery (kurgan plans redrawn from Epimakhov, 2005: 10, 79).

The KA-5 (MBA), Bestamak (MBA) and Lisakovsk (LBA) datasets exhibited a wide range of δ13C and δ15N values for both humans and herbivores (Figs. 5 and 6 & Table 8). This diversity in isotopic signals may be evident for a variety of reasons. For example, the range of values may be associated with a broad spectrum of C3 and C4 plant diversity in the ancient site biome or herbivore grazing patterns that included more diverse environmental niche areas in the microregion around the sampled sites. Herders also may have chosen to graze animals in niche areas due to recognized territorial boundaries between settlements and concomitant patterns of mobility. Importantly, data from Bolshekaragansky represents humans with lower δ15N values that are more closely associated with δ15N values of the sampled domestic herbivores (Fig. 6). When the archaeological evidence from associated settlement sites is considered, Bolshekaragansky, Bestamak, Lisakovsk and KA-5 have been assumed to represent populations that shared similar forms of pastoral subsistence economies with significant dietary reliance upon domesticated herbivore meat and milk. Human diets have δ13C values closely related to those of local herbivores in terms of the slope of the trendline and range of values (Fig. 6). Comparatively, the cemetery of Bolshekaragansky (associated with the Arkaim settlement) reflects individuals with trend lines closer to those of cattle and caprines and may indicate a stronger reliance on subsistence products from these species with less use of wild riverine and terrestrial resources. The site of Čiča is significantly different with elevated human δ15N isotopic values and depleted δ13C values indicative of a subsistence regime more closely associated with the consumption of freshwater resources, such as fish. The stable isotopic data in this instance is strongly supported by zooarchaeological evidence recovered from the Čiča settlement and also is indicative of significant diachronic changes from the LBA phases through the Iron Age (Fig. 6).

kamenniy-ambar-isotopic-chicha-lisakovsk-bestamark
Regional analysis and comparison of stable isotope results from humans (adults) and animals recovered from MBA and LBA cemeteries in the Southern Urals (Kamennyi Ambar 5 & Bolshekaragansky) northwestern Kazakhstan (Liskovsk & Bestamak) and southwestern Siberia (Čiča).

Conclusion

(…) The isotopic results from KA-5, and recent botanical and archaeological studies from the Kamennyi Ambar settlement, have not produced any evidence for the production or use of domesticated cereals. While this does not definitively answer the question as to whether Sintashta populations engaged in agriculture and/or utilized agricultural products, it does call into serious question the ubiquity of such practices across the region and correlates well with recent archaeological, bioarchaeological, and isotopic studies of human and animal remains from the Southwestern Urals region and Samara Basin (Anthony et al., 2016; Schulting and Richards, 2016). The results substantiate a broader spectrum subsistence diet that in addition to the use of domesticated animal products also incorporated wild flora, wild fauna and fish species. These findings further demonstrate the need to draw on multiple methods and datasets for the reconstruction of late prehistoric subsistence economies in the Eurasian steppes. When possible, this should include datasets from both settlements and associated cemeteries.

Variability in subsistence practices in the central steppes region has been highlighted by other scholars and appears to be strongly correlated with local environmental conditions and adaptations. More comprehensive isotopic studies of human, animal and fish remains are of fundamental importance to achieve more robust and empirically substantiated reconstructions of local biomes and to aid the refinement of regional and micro-regional economic subsistence models. This will allow for a fuller understanding of key diachronic shifts within dietary trends and highlight regional variation of such practices. Ultimately, this will more effectively index the diverse social and environmental variables that contributed to late prehistoric lifeways and the economic strategies employed by these early steppe communities.

Social organization of Sintashta-Petrovka

Interesting to remember now the recent article by Chechushkov et al. (2018) about the social stratificaton in Sintashta-Petrovka, and how it must have caused the long-lasting, peaceful admixture process that led to the known almost full replacement of R1b-L23 (mostly R1b-Z2103) by R1a-Z645 (mostly R1a-Z93) subclades in the North Caspian steppe, coinciding with the formation of the Proto-Indo-Iranian community and language (read my thoughts on this after Damgaard et al. 2018).

Here is another relevant excerpt from Chechushkov et al. (2018), translated from Russian:

settlement-kamenniy-ambar
The map of the settlement of Kamennyi Ambar with excavations, soil cores, and test pits. Legend: a — cuts of the sides of ravines; b — test pits of 2015—2017; c — test pits of 2004; d — soil-science samples with a cultural layer; e — soil-science samples without cultural layer; f — borders of archaeological sites (interpretation of the plan of magnetic anomalies); g — boundaries of excavated structures (1, 2, 4, 5, 7 — Sintashta-Petrovka culture; 3, 6 — Srubnaya-Alakul’ culture).

The analysis suggests that the Sintashta-Petrovka societies had a certain degree of social stratification, expressed both in selective funeral rituals and in the significant difference in lifestyle between the elite and the immediate producers of the product. The data obtained during the field study suggest that the elite lived within the fortifications, while a part of the population was outside their borders, on seasonal sites, and also in stationary non-fortified settlements. Probably, traces of winter settlements can be found near the walls, while the search for summer ones is a task of a separate study. From our point of view, the elite of the early complex societies of the Bronze Age of the Eurasian steppe originated as a response to environmental challenges that created risks for cattle farming. The need to adapt the team to the harsh and changing climatic conditions created a precedent in which the settled collectives of pastoralists – hunter-gatherers could afford the content and magnificent posthumous celebration of people and their families who were not engaged in the production or extraction of an immediate product. In turn, representatives of this social group directed their efforts to the adoption of socially significant decisions, the organization of collective labor in the construction of settlement-shelters and risked their lives, acting as military leaders and fighters.

Thus, in Bronze Age steppe societies, the formation, development and decline of social complexity are directly related to the intensity of pastoralism and the development of new territories, where collectives had to survive in part a new ecological niche. At the same time, some members of the collective took upon themselves the organization of the collective’s life, receiving in return a privileged status. As soon as the conditions of the environment and management changed, the need for such functions was virtually eliminated, as a result of which the privileged members of society dissolved into the general mass, having lost their lifetime status and the right to be allocated posthumously.

Also interesting for the MLBA haplogroup bottleneck in the region is the paper by Judd et al. (2017) about fast life history in Early Indo-Iranian territories.

On the arrival of haplogroup N1c1-L392

Regarding the special position of the Chicha-1 samples in the change of diet and economy during the Iron Age, it is by now well known that haplogroup N must have arrived quite late to North-East Europe, and possibly not linked with the expansion of Siberian ancestry – or linked only with some waves of Siberian ancestry in the region, but not all of them. See Lamnidis et al. (2018) for more on this.

Also, the high prevalence of haplogroup N among Fennic and Siberian (Samoyedic) peoples is not related: while the latter reflects probably the native (Palaeo-Siberian) population that acquired their Uralic branch during the MLBA expansions associated with Corded Ware groups, the former points to the expansion of Fennic peoples into Saamic territory (i.e. after the Fenno-Saamic split) as the most likely period of expansion of N1c1-L392 subclades (see known recent bottlenecks among Finns, and on Proto-Finnic dialectalization).

Probably related to these late incomers are the ancient DNA samples from the Sargat culture during the Iron Age, which show the arrival of N subclades in the region, replacing most – but not all – R1a lineages (see Pilipenko et al. (2017)). Regarding the site of Chicha-1, the following are relevant excerpts about the cultural situation that could have allowed for such stepped, diachronic admixture events in Northern Eurasia, from the paper Stages in the settlement history of Chicha-1: The Results of ceramic analysis, by Molodin et al. (2008):

The stratigraphic data allows us to make the following inference: originally, the settlement was inhabited by people bearing the Late Irmen culture. Later, the people of the Baraba trend of the Suzgun culture arrived at the site (Molodin, Chemyakina, 1984: 40–62). The Baraba-Suzgun pottery demonstrates features similar to what has been reported from the sites of the transitional Bronze to Iron Age culture in the pre-taiga and taiga zones in the Irtysh basin (Potemkina, Korochkova, Stefanov, 1995; Polevodov, 2003). The major morphological types are slightly and well-profiled pots with a short throat. (…)

chicha-irmen-tagar-baraba-forest-siberian
Map showing the location of Chicha-1.

During the following stage of development of the site, the Chicha population increased with people who practiced cultures others than those noted in earlier collections. The ceramic materials from layer 5 provide data on possible relationships. In addition to migrants from northwestern regions practicing the Suzgun culture, there were people bearing the Krasnoozerka culture. Available data also suggests that people from the northern taiga region with the Atlym culture visited the site.

However, people from the west and southwest represent the greatest migration to the region under study. In all likelihood they moved from the northern forest-steppe zone of modern Kazakhstan and practiced the Berlik culture. The spatial distribution analysis of the Chicha-1 site suggests that the Berlik population was rather large. The Berlik people formed a single settlement with the indigenous Late Irmen people and apparently waged certain common economic activities, but preserved their own ethnic and cultural specificity (Molodin, Parzinger, 2006: 49–55). Judging by the data on the chronological sequence of deposited artifacts, migration took place roughly synchronously, hence Chicha-1 became a real cultural and economic center.

(…) In sum, the noted distribution of ceramics over the culture-bearing horizons suggests that beginning with layer 5, traditions of ceramic manufacture described above were practiced, hence the relevant population inhabited the site. Apparently, there were two predominant traditions: the local Late Irmen cultural tradition and the Berlik tradition, which was brought by the immigrants. The Late Irmen people mostly populated the citadel, while the Berlik immigrants inhabited the areas to the east and the north of the citadel.

The stratigraphic data also suggest that the Early Sargat ceramics emerged at the site likely as a part of the Late Irmen tradition (…) Early Sargat ceramics is apparently linked with the Late Irmen tradition. Artifacts associated with the Sargat culture proper have been found in several areas of Chicha-1 (e.g., in excavation area 16). However, the Sargat people appeared at the site after it had been abandoned by its previous inhabitants, and had eventually become completely desolated. This happened no earlier than the 6th cent. BC, possibly in the 5th cent. BC (in fact, the radiocarbon dates for that horizon are close to the turn of the Christian era).

Related

BMAC: long term interaction between agricultural communities and steppe pastoralists in Central Asia

indo-european-indo-iranian-migrations

Interesting new paper Mixing metaphors: sedentary-mobile interactions and local-global connections in prehistoric Turkmenistan, by Rouse & Cerasetti, Antiquity (2018) 92:674-689.

Relevant excerpts (emphasis mine):

The Murghab alluvial fan in southern Turkmenistan witnessed some of the earliest encounters between sedentary farmers and mobile pastoralists from different cultural spheres. During the late third and early second millennia BC, the Murghab was home to the Oxus civilisation and formed a central node in regional exchange networks (Possehl 2005; Kohl 2007). The Oxus civilisation (or the Bactria-Margiana Archaeological Complex) relied on intensive agriculture to support a hierarchical society and specialised craft production of metal and precious stone objects for prestige display and long-distance exchange (Sarianidi 1981; Hiebert 1994). By c. 1800 BC (the local Late Bronze Age), the internal coherence of the Oxus civilisation began to break down, along with the inter-regional exchange networks; the settlement structure of the Murghab shifted from a tiered system of urban centres, villages and hamlets, to a more dispersed pattern of smaller-scale agricultural settlements (Salvatori 2008). Contemporaneous evidence for small campsites (with a distinct ceramic tradition) suggests an influx of mobile pastoralists from the Central Eurasian Steppe and foothills (Cerasetti 1998; Masson 2002; Cattani et al. 2008). This striking combination of the sites and material cultures of both late Oxus farmers and ‘steppe’ pastoralists spans more than 500 years of Murghab prehistory (Salvatori 2008; Rouse & Cerasetti 2017).

The mixed farmer-pastoralist archaeological record of the Murghab has influenced competing interpretations of Later Bronze Age socio-political and economic relationships. Some scholars argue that the ‘collapse’ of the Oxus civilisation was at least partly due to the hostile incursions of nomads (Marushchenko 1956; Kuz’mina&Lyapin 1984; Vinogradova & Kuz’mina 1996). Others suggest that pastoralists took advantage of the Murghab’s crumbling power structure by moving into the area, but occupying only marginal, agriculturally unsuitable zones (P’yankova 1993), or merging with the late Oxus farming populations (Masson 2002). These models broadly follow ‘trade or raid’ paradigms of farmer-pastoralist interaction, whereby the perceived shortages of pastoralist communities force them to rely on agriculturalists for subsistence, material and cultural inputs (Kroeber 1947; Ferdinand 2003; Potts 2014). Such models may explain certain cases of Near Eastern pastoral economic specialisation, or historical contact scenarios between Eurasian steppe and agricultural communities on China’s northern frontier (Lattimore 1979; Barfield 2001; Alizadeh 2009; Khazanov 2009). Near Eastern and Eurasian interaction paradigms, however, fit increasingly poorly with the archaeological evidence for early farmer-pastoralist encounters in southern Central Asia.

We present data from four Murghab pastoralist campsites dating to the third to second millennia BC, restricting our discussion to the materials and practices employed by Oxus-period pastoralists to navigate shifting social, political and economic networks. Our aim is to highlight how variable strategies broadly identified under the rubric of ‘agropastoralism’ can be teased apart to recognise mechanisms of social boundary-making. Individually, these four sites present chronologically and locally distinct snapshots of farmer-pastoralist interactions across different realms of exchange (e.g. subsistence, technology and ideology); they provide examples of how pastoralists and farmers mutually participated in each other’s material and social norms. Together, these sites reveal how varied farmer-pastoralist engagement with technology and material culture did not lead inevitably to the assimilation of the two groups; rather, they worked consciously within existing systems of cultural practice to maintain distinct ‘farmer’ and ‘pastoralist’ identities, potentially over a 900-year period.

oxus-bmac-pastoralism
Region of Central Asia as discussed in this article. Areas traditionally identified with farming-dependent Oxus communities and non-Oxus mobile pastoralists are shown, acknowledging that in both areas mixed agropastoral practices have occurred in the past and present.

Conclusions

(…)First, the results indicate a cultural model of ‘being’ a pastoralist that was maintained actively over hundreds of years, in part by its material difference from that of local farmers. Second, the variability of materials, technologies and practices shared at these campsites suggests that no hegemonic power controlled trade relationships or regulated economic dependency between Oxus farmers and non-Oxus mobile pastoralists in the Murghab. Indeed, current data indicate that pastoralist occupation in the Murghab intensified during the waning of Oxus political centralisation, suggesting that the loosening of state-level structures provided the opportunity for intercultural interactions, rather than interactions being promoted or facilitated from the top. Finally, in the removal of broad-brush narratives that polarise ‘the steppe’ and ‘the sown’, and the integration of evidence suggesting that mobile pastoralists influenced the crop systems of farmers in southern Central Asia (Spengler et al. 2014b), these four sites allow us to recognise the means by which farmers and pastoralists re-shaped cultural institutions while reinforcing the meaningfulness of the associated social categories. Current work in the Murghab complements detailed studies of pastoralists in other Eurasian contexts (e.g. Frachetti 2008; Rogers 2012; Honeychurch 2015) in beginning to unravel simplistic notions of broad cross-cultural exchanges in Eurasian prehistory and the political entities traditionally seen as directing them.

The whole article is very interesting, and the four sites studied and their relevance for the said interactions are described in detail, and in chronological order. If you have the opportunity, read it.

I found it interesting that the article mentions the traditional scholarly opposition of agriculturalists vs. pastoralists (‘civilised/barbarian’, ‘state/tribe’ and ‘centre/periphery’) as an idea of Eurasian origin, and having deep ‘Western’ roots. Reading what many OIT (or anti-AIT, as they like to call themselves) supporters write, it seems to me as though they have entirely accepted and in fact are eager to promote this ‘Western’ narrative from the mid-20th century…

Steppe MLBA

This is what Narasimhan et al. (2018) had to say about the BMAC – Steppe pastoralists interaction:

We document a southward spread of genetic ancestry from the Eurasian Steppe, correlating with the archaeologically known expansion of pastoralist sites from the Steppe to Turan in the Middle Bronze Age (2300-1500 BCE). These Steppe communities mixed genetically with peoples of the Bactria Margiana Archaeological Complex (BMAC) whom they encountered in Turan (primarily descendants of earlier agriculturalists of Iran), but there is no evidence that the main BMAC population contributed genetically to later South Asians. Instead, Steppe communities integrated farther south throughout the 2nd millennium BCE, and we show that they mixed with a more southern population that we document at multiple sites as outlier individuals exhibiting a distinctive mixture of ancestry related to Iranian agriculturalists and South Asian hunter-gathers.

yamna-steppe-emba-mlba-cloud
Narasimhan et al. (2018): “Modeling results.(A) Admixture events originating from 7 “Distal” populations leading 538 to the formation of the modern Indian cloud shown geographically. Clines or 2-way mixtures of 539 ancestry are shown in rectangles, and clouds (3-way mixtures) are shown in ellipses.

(…) The absence in the BMAC cluster of the Steppe_EMBA ancestry that is ubiquitous in South Asia today—along with qpAdm analyses that rule out BMAC as a substantial source of ancestry in South Asia (Fig. 3A)—suggests that while the BMAC was affected by the same demographic forces that later impacted South Asia (the southward movement of Middle to Late Bronze Age Steppe pastoralists described in the next section), it was also bypassed by members of these groups who hardly mixed with BMAC people and instead mixed with peoples further south. In fact, the data suggest that instead of the main BMAC population having a demographic impact on South Asia, there was a larger effect of gene flow in the reverse direction, as the main BMAC genetic cluster is slightly different from the preceding Turan populations in harboring ~5% of their ancestry from the AASI.

(…)between 2100-1700 BCE, we observe BMAC outliers from three sites with Steppe_EMBA ancestry in the admixed form typically carried by the later Middle to Late Bronze Age Steppe groups (Steppe_MLBA). This documents a southward movement of Steppe ancestry through this region that only began to have a major impact around the turn of the 2nd millennium BCE.

Related