Archaeology, Genetics, and Language in the Steppes: A Comment on Bomhard David W. Anthony Hartwick College Bomhard’s hypothesis is that PIE was the result of interference between a substrate related to Northwest Caucasian and a dominant language related to Uralic (pre-Uralic?) that absorbed Caucasus-like elements in phonology, morphology, and lexicon. That kind of interference would imply a long period of widespread bilingualism among the pre-Uralic speakers. The shared lexical cognates that Bomhard lists include kinship terms such as ‘daughter-in-law’, suggesting the occurrence of at least occasional formal intermarriage between the two language communities. I have been asked to outline how this hypothesis might correlate with genetic and archaeological evidence ‘on the ground’ in the Pontic-Caspian-North Caucasus region. Much of my assessment is based on research that has been posted on the public server bioarxiv but is not yet formally published. I accept Mallory’s reading of the current consensus that the Yamnaya expansion, beginning about 3000 BC into both Europe and Asia from the Pontic-Caspian steppes, represented the expansion of late PIE languages (after the separation of Anatolian). Putting aside the questions of how and why that expansion occurred, my topic is the formation and origin of the Yamnaya mating network, as a genetic phenomenon; and secondarily of the Yamnaya culture, beginning about 3300 BC within the Pontic-Caspian steppes, as an archaeological phenomenon. I also assess how pre-Yamnaya genetic and archaeological patterns of interaction might correlate with Bomhard’s hypothesis for early PIE origins. Mating networks and IE origins I should note two important facts about ancient DNA (aDNA) studies. First is that large-scale studies involving hundreds of prehistoric individuals analyzed at the whole-genome scale only became methodologically feasible recently, and the first results were published only in 2015 (Haak et al. 2015; Allentoft et al. 2015). So the evidentiary basis under our feet is recently formed Volume 47, Number 1 & 2, Spring/Summer 2019 2 David W. Anthony and is changing as we read. Second, sample sizes for aDNA studies should not be thought of in the same way that we think of adequate sample sizes for pottery sherds. Because each person carries a genetic record of descent from hundreds of recognizable individual ancestors (up to 10 generations back) and many more broadly defined ancestral groups, the whole genomes of a few individuals can reveal the histories of substantial populations. Applied to large samples of individuals, statistical characterizations of genetic relationships can identify migrations, reveal their demographic structure, and describe ancient mating networks – a new category of measurable human relationships. These topics previously were debated or unknowable. We are just beginning to formulate our interpretations in light of this new information. I use the term ‘mating network’ for genetically linked populations that shared a distinctive group of genetic traits as determined by ancient DNA (aDNA) analyzed across whole genomes, such that individuals from that chronological period and part of the world can confidently be assigned to one mating network rather than another. The aDNA groups were defined by geneticists since 2015, but I have not seen a standardized label for referring to them. ‘Mating network’ describes the underlying behavior. Mating networks could be (but were not always) very large in scale, much larger than local dialects or material culture groups. They could have been the product of a combination of different mating behaviors (rape, concubinage, informal relations, marriage) with the qualification that they were engaged in often enough to contribute visibly to the genetic admixture of the linked populations. In most cases sustained genetic mixing sufficient to produce broad genetic affinity within a mating network would have been associated with sustained formal marriages between linked populations within the network, possibly augmented by sustained slave raiding and concubinage between some linked populations, a practice found among many tribal societies (Cameron 2016). One important mating network that contributed to Yamnaya genetic ancestry was the Eastern Hunter-Gatherers (EHG). EHG averaged about 50% of the ancestry of the Yamnaya populations that have been studied to date (Haak et al. 2015; Allentoft et al. 2015; Damgaard et al. 2018). The EHG The Journal of Indo-European Studies Archaeology, Genetics, and Language in the Steppes 3 genetic type was first defined in Haak et al. (2015: 208) on the basis of two genetically similar Mesolithic/Neolithic individuals from Lebyazhinka IV near Samara and Oleni Ostrov in Lake Onega near the Baltic in Russia. Eastern Hunter-Gatherers ranged from the Urals to the Baltic and moved down the river valleys into the steppes north of the Black and Caspian Seas. The EHG mating network was a Holocene, western regional survival of a much larger Upper Paleolithic mating network called Ancient North Eurasian (ANE), first recognized in Upper Paleolithic humans who lived near Lake Baikal, at a site called Mal’ta about 24,000 years ago and at another site, Aftonova Gora, about 16,000 years ago (Raghavan et al. 2014). Interconnected mating networks distributed variants of the ANE genetic type across northern Eurasia during the Upper Paleolithic, when mammoths and mammoth-hunters roamed across Siberia (i
ánková et al. 2014). These networks extended into North America on the east (where ANE averages 15-40% of Native American genetic ancestry) and into the Pontic–Caspian steppes and the Baltic forests on the west. Within the EHG mating network ANE averaged about 70% of the ancestry of hunter-fishers of the sixth millennium BC (Haak et al. 2015). This high level of ANE ancestry distinguished the EHG from the Western Hunter-Gatherers (WHG) who spread across western and Mediterranean Europe after about 15,000 years ago, possibly from a glacial refuge in southeastern Europe, who showed very little affinity with the ANE. Like the EHG mating network, the WHG mating network was enormous in scale, as it extended from Britain (so-called Cheddar Man) to the lower Danube valley (the Iron Gates region) by 6000 BC. The frontier between WHG and EHG populations emerged after 7000 BC as a broad zone of genetic admixture from the western Baltic down to what is today Romania and western Ukraine, between the lower Danube valley (the Mesolithic cemeteries in the Iron Gates contained a majority of WHG individuals) and the forests west of the Dnieper River. Mesolithic and Early Neolithic cemeteries in the Dnieper valley contained individuals with largely EHG ancestry, but admixed consistently and continuing through millennia (7000-5000 BC) with recurring WHG mates (Mathieson et al 2018). This EHG/WHG admixture continued Volume 47, Number 1 & 2, Spring/Summer 2019 4 David W. Anthony into the Eneolithic, at Dereivka-1 and Vil’nyanka, which continued to show a population with y-chromosome EHG/WHG ancestries through the sixth and fifth millennia BC. Figure 1. Sites mentioned in the text. East of the Dnieper, EHG ancestry un-admixed with WHG was dominant among the hunters and fishers who occupied the river valleys that ran south through the Pontic–Caspian steppes into the Black and Caspian Seas (the Donets, Don, Volga, and Ural rivers) before the Yamnaya period (before 3300-2600 BC in the Pontic-Caspian steppes). EHG averaged about 50% of the ancestry of the Yamnaya populations. The other half of Yamnaya genetic ancestry was Caucasus Hunter-Gatherer (CHG). The CHG mating network occupied the Caucasus Mountains, parts of eastern Anatolia, and the western Iranian plateau. How and when a CHG population entered the steppes and came to contribute half of Yamnaya genetic ancestry is an open question, and crucial for Bomhard’s hypothesis. The archaeological sites where the CHG type was first identified The Journal of Indo-European Studies Archaeology, Genetics, and Language in the Steppes 5 were a late Upper Paleolithic burial at Satsurblia in western Georgia dated 11,200 BC and a Mesolithic grave at Kotias Cave, also in western Georgia, dated 7500–7800 BC (Jones et al. 2015). These two Paleolithic and Mesolithic individuals were quite similar genetically to a Mesolithic forager dated 8600–9100 BC from level IIIb at Hotu Cave in northern Iran on the southeastern Caspian coast, located 1300 km southeast of the Georgian sites and in a very different ecological setting (Lazaridis et al. 2016). People with primarily CHG ancestry also lived 750 km southwest of Hotu Cave, at Early Neolithic Ganj Dareh near Kermanshah, dated 7000 BC (Lazaridis et al. 2016). Because people with CHG ancestry lived on the northern (Hotu Cave) and southern (Ganj Dareh) sides of the western Iranian plateau, a similar CHG population probably was distributed across western Iran and the Caucasus at the beginning of the Neolithic. A signal of diluted CHG ancestry appeared in some Neolithic individuals at Tepecik-Çiftlik in central Anatolia dated 6500 BC. Tepecik-Çiftlik can be seen as the far western edge, the tail of the curve of the CHG mating network, around 6500 BC (Kliniç et al. 2017). Most Yamnaya genomes studied to date exhibit admixed EHG & CHG ancestry with each in robust proportions, often with CHG ancestry higher than 50% (Wang et al. 2018: Figure 2c). This north-south genetic mixture would seem to support Bomhard’s linguistic analysis of PIE as a north-south admixture between a pre-Uralic language, easily placed in EHG territory; and a Northwest Caucasian language, easily assigned to CHG territory. To be more specific, the Maikop culture (3600-3000 BC) is regarded in many scenarios as the likely source of the CHG that mixed with steppe EHG mating networks to create the Yamnaya genetic synthesis. In this case, the language of the Maikop culture could represent the ancestor of the Anatolian branch, and the Maikop culture could be the material correlate of the early PIE homeland, while Yamnaya represented the spread of PIE across the steppes and then beyond, from the late PIE homeland (Reich 2018: 107–109, 120; Kristiansen et al. 2018). But if we look more closely at the genetics and the cultural evidence from archaeology, this simple match between two two-way admixtures, one linguistic and the other genetic, Volume 47, Number 1 & 2, Spring/Summer 2019 6 David W. Anthony becomes complicated, like everything connected with IndoEuropean origins. How to make a Yamnaya cocktail: a pinch of Anatolian Farmer, but from where? A recently discovered complicating factor is a component of previously undetected Anatolian Farmer ancestry, recently found in Yamnaya genomes by Wang et al. (2018). They reported that the Anatolian Farmer component varied from about 10 to 18% of ancestral DNA across three sub-regions of the Yamnaya culture in the steppes (although this component is absent from the color-coded bar-graphs of Yamnaya ancestry in Wang et al. 2018, figure 2c). The proportion of farmer ancestry was lowest in the Volga-Ural steppes and highest in the Ukrainian steppes, with the North Caucasus steppes between, but the difference between the sub-regions was not statistically significant. Where did this ancestry come from? The Anatolian Farmer mating network evolved in western Anatolia during the Neolithic, ca. 7500-5500 BC. A subpopulation or populations migrated first to Greece and Crete, then through temperate Europe and the Mediterranean, establishing the greatly expanded mating network that characterized the first Neolithic farmers of Europe (Haak et al. 2015; Mathieson et al 2018). Anatolian Farmers of course remained living in western Anatolia, and after about 5000 BC their mating networks expanded eastward to include mates in Transcaucasia (Areni-1 Chalcolithic, about 4200-4000 BC) and western Iran (Seh Gabi Chalcolithic, about 4800-4600 BC). The Anatolian Farmer ancestry recently discovered in Yamnaya genomes could have entered steppe mating networks through the Caucasus or through southeastern Europe. Its geographic source is an important question that also affects the source of the CHG in Yamnaya. When the first Yamnaya whole genomes were published (Haak et al. 2015; Allentoft et al 2015) two independent labs concluded that the Yamnaya individuals from the Volga and North Caucasus steppes had no Anatolian Farmer ancestry. But with larger and older samples from the Caucasus and Iran, and much larger and older samples from the steppes, the reference groups used to calculate genetic distances have changed values The Journal of Indo-European Studies Archaeology, Genetics, and Language in the Steppes 7 in both regions, while the networks themselves have remained. In conversation, colleagues in David Reich’s lab at Harvard agreed with Wang et al. (2018) that the Yamnaya mating network in the Pontic-Caspian steppes probably contained some Anatolian Farmer ancestry. Wang et al. (2018) concluded that the specific kind of Anatolian Farmers who mated into the Yamnaya mating network were European farmers, and not from the Caucasus. They pointed out that samples from the steppe-adjacent late Tripol’ye or Globular Amphorae cultures in western Ukraine, among others, had close to the right mix of 80% Anatolian Farmer with 20% WHG, an admixture that appeared across western and central Europe (but not in the Caucasus) in the fourth millennium BC, reflecting a rebound in the old indigenous WHG population. People with approximately this ancestry, 80/20 Anatolian Farmer/WHG, mated into the Yamnaya population occasionally, accounting for 10-18% of Yamnaya ancestry. It was the faint trace of WHG, perhaps 3% of whole Yamnaya genomes, that identified this admixture as coming from Europe, not the Caucasus, according to Wang et al. (2018). Colleagues in David Reich’s lab commented that this small fraction of WHG ancestry could have come from many different geographic places and populations. But if Wang et al. 2018 are correct, if the Anatolian Farmer element in Yamnaya came from mating between steppe people and late Tripol’ye or Globular Amphorae farmers, then the CHG genetic element in Yamnaya, more than half of Yamnaya ancestry, could not have been heavily admixed with additional Anatolian Farmer ancestry. Yamnaya had a minor component of Anatolian Farmer ancestry, 10-18%. If most of it came from Europe then the CHG that was so prominent in Yamnaya ancestry must have been a relatively un-admixed variety of CHG, with a low percentage of Anatolian Farmer ancestry. This un-admixed kind of CHG disappeared after about 5000 BC in the Caucasus and northwestern Iran, according to Wang et al. (2018) combined with Lazaridis et al. (2016) and the forthcoming Naramsimhan et al. (2018 posted on bioarxiv). After about 5000 BC Anatolian Farmer ancestry spread east through eastern Anatolia and Transcaucasia (Areni-1, Armenia) into Iran (Seh Gabi) while CHG ancestry spread westward into Volume 47, Number 1 & 2, Spring/Summer 2019 8 David W. Anthony Anatolia and the Levant. All tested individuals dated after 5000 BC in the Caucasus and western Iran showed CHG & Anatolian Farmer admixture on a cline across Iran with greater Anatolian Farmer ancestry in western Iran and the Caucasus and less to none in eastern Iran (Narasimhan et al. 2018). After this mixing of populations happened, the un-admixed type of early CHG ancestry probably survived only in small isolated populations. According to Wang et al. (2018), the Maikop population was descended from the Eneolithic farmers who first colonized the north side of the North Caucasus ridge. They came from the south, probably from western Georgia and Abkhazia, today home to Northwest Caucasian languages. They are called the Darkveti-Meshoko culture by Trifonov in Wang et al. (2018), and are the ideal archaeological candidate for the founders of the Northwest Caucasian language family. From sites like Darkveti Cave on the Georgia/Abkhazia side of the mountains, a region where northwest Caucasian languages are spoken today, they migrated over the North Caucasus ridge into the upper Belaya river valley, the other principal region where Northwest Caucasian languages are spoken today. Their earliest known arrival is dated about 4600-4500 BC at Unakozovskaya Cave in the upper Belaya River valley (Wang et al. 2018). Rapidly after that a handful of walled agricultural settlements, including Meshoko, spread down the Belaya valley to the edge of the steppes (Svobodnoe) before 4000 BC. They established farming (wheat) and domesticated animal herding (cattle, pigs, sheep & goats) in the northwest Caucasus region, and also founded the genetic mating network that later would characterize the people of the Maikop culture, their direct descendants. The great Maikop tumulus, the astonishingly rich grave that marked the sudden transition from low to very steep hierarchy dated about 3600 BC, was located in the longestsettled part of the North Caucasus, in the Belaya River valley. As Mallory noted in the Introduction, the Maikop culture (36003000 BC) is regarded in many scenarios as the likely source of the CHG in Yamnaya genomes. Its oldest phase was in a region where farming first began in the North Caucasus, and where Northwest Caucasian languages are spoken today, so a Maikop origin for Yamnaya CHG would strongly support Bomhard’s hypothesis. The Journal of Indo-European Studies Archaeology, Genetics, and Language in the Steppes 9 But Wang et al. (2018) complicated that picture. From the first Eneolithic farmers who appeared at Unakozovskaya in the northwest Caucasus about 4600 BC to the Maikop culture, all of them had significant CHG ancestry, on a cline of 28-60%. But the other 72-40% of their ancestry can be modeled as an admixture of Anatolian Farmer (the great majority), Levantine Neolithic, and Ganj-Dareh-type Iranian Farmer ancestries. The Anatolian Farmer component is bar-graphed as 30-40% of the Eneolithic farmers’ ancestry (Wang et al. 2018: Figure 2c). Similar percentages characterized the Maikop population. This mixture was too rich in Anatolian Farmer genes to have contributed much to the Yamnaya gene pool, which had only 10-18% Anatolian Farmer ancestry, and most of that arguably derived from the west, from Globular Amphorae and late Tripol’ye populations. If Wang et al (2018: line 591 on bioarxiv) are correct that Yamnaya and all later steppe populations “deviate from the [Eneolithic steppe population’s] EHG/CHG towards European populations in the West” then Maikop is left to play only a small role in Yamnaya ancestry, less than Europe. Also, the Y-chromosome haplogroups of the Eneolithic Meshoko and Maikop men were typical AnatolianIranian Neolithic haplogroups (L, J2, and G2) unlike the paternal haplogroups of the steppes. Yamnaya men were almost exclusively R1b, and pre-Yamnaya Eneolithic Volga-CaspianCaucasus steppe men were principally R1b, with a significant Q1a minority. Maikop men did not father a significant number of Yamnaya males. If there was any Maikop gene flow into Yamnaya, it could have been through a small number of Maikop females whose 30-40% Anatolian Farmer ancestry was diluted in their descendants, and whose skeletons have not yet been found or analyzed. This partial description of the genetic data, if it stands, suggests that Maikop was not the source of most of the CHG that amounts to half of Yamnaya ancestry. This is because CHG was already in the steppes long before Maikop, and it was in an unadmixed form. This older introduction of CHG into the steppes is less compatible with the Maikop-NWCaucasian-Yamnaya-PIE connection, but it is what is indicated by the emerging genetic data. Volume 47, Number 1 & 2, Spring/Summer 2019 10 David W. Anthony CHG in the steppes 6000-4000 BC The variety of CHG that constituted more than half of Yamnaya ancestry could have been the Mesolithic/Early Neolithic variety, like Hotu Cave or Kotias Cave, not yet admixed with Anatolian Farmer ancestry. If the CHG element in Yamnaya came from a non-admixed CHG population of this kind, they could have walked into the steppes from northwestern Iran/Azerbaijan at any time before about 5000 BC—before admixture with Anatolian Farmers began. The easiest path into the steppes from the Caucasus skirts the eastern end of the North Caucasus ridge, where there is a level plain between the mountains and the Caspian Sea. No such plain exists at the western end of the mountain range, where mountains plunge into the Black Sea. Of course this eastern origin for CHG is not so easily matched with western Caucasian languages. But archaeologists have noticed similarities in the dates of adoption of ceramics and agriculture in sites scattered around the southern end of the Caspian Sea (Gorgan plain) and in the southeastern Caucasus region (Kamiltepe and others in Azerbaijan). Helwing (2012: 348) wrote of a “seaborne interaction sphere” that connected these regions already in the EpiPaleolithic/Mesolithic that continued into the Neolithic. Neolithic Kamiltepe at about 5500 BC had painted ceramics and mudbrick architecture that certainly never appeared in the steppes, but a Caucasian CHG population from an older phase, before 5500 BC, could have migrated from the south Caspian to the north Caspian, which would put them in the steppes. Hunter-fisher groups from the demonstrated south Caspian CHG network could have been attracted to the north Caspian by the rich resources of the Volga estuary, the largest river estuary in temperate Eurasia (Figure 2). Here were streams full of sturgeons, with herds of wild onagers, horses, and saiga antelope running through the grasslands beside the river, and broad marshes filled with birds and fish. Natural saltpans near the coast produced a valued trade item. We don’t have published aDNA samples from lower Volga-Caspian steppe archaeological sites because graves were few and scattered, but we do have archaeology, none of it in English, The Journal of Indo-European Studies Archaeology, Genetics, and Language in the Steppes 11 although I summarized what I knew 10 years ago (Anthony 2007: 188-192). Figure 2. Eneolithic sites on the middle and lower Volga, from Morgunova 2015. Along the banks of the lower Volga many excavated hunting-fishing camp sites are dated 6200-4500 BC. They could be the source of CHG ancestry in the steppes. At about 6200 BC, when these camps were first established at Kair Shak III and Varfolomievka (42 and 28 on Figure 2), they hunted primarily saiga antelope around Dzhangar, south of the lower Volga, and almost exclusively onagers in the drier desertsteppes at Kair Shak, north of the lower Volga. Farther north at the lower/middle Volga ecotone, at sites such as Varfolomievka and Oroshaemoe hunter-fishers who made pottery similar to that at Kair-Shak hunted onagers and saiga antelope in the desert-steppe, horses in the steppe, and aurochs in the riverine forests. Finally, in the Volga steppes north of Saratov and near Samara, hunter-fishers who made a different kind of pottery (Samara type) and hunted wild horses and red deer definitely were EHG. A Samara hunter-gatherer of this era buried at Lebyazhinka IV, dated 5600-5500 BC, was one of the first named examples of the EHG genetic type (Haak et al. 2015). This individual, like others from the same region, had no or very Volume 47, Number 1 & 2, Spring/Summer 2019 12 David W. Anthony little CHG ancestry. The CHG mating network had not yet reached Samara by 5500 BC. But before 4500 BC, CHG ancestry appeared among the EHG hunter-fishers in the middle Volga steppes from Samara to Saratov, at the same time that domesticated cattle and sheepgoats appeared. The Reich lab now has whole-genome aDNA data from more than 30 individuals from three Eneolithic cemeteries in the Volga steppes between the cities of Saratov and Samara (Khlopkov Bugor, Khvalynsk, and Ekaterinovka), all dated around the middle of the fifth millennium BC. Many dates from human bone are older, even before 5000 BC, but they are affected by strong reservoir effects, derived from a diet rich in fish, making them appear too old (Shishlina et al 2009), so the dates I use here accord with published and unpublished dates from a few dated animal bones (not fish-eaters) in graves. Only three individuals from Khvalynsk are published, and they were first published in a report that did not mention the site in the text (Mathieson et al. 2015), so they went largely unnoticed. Nevertheless, they are crucial for understanding the evolution of the Yamnaya mating network in the steppes. They were mentioned briefly in Damgaard et al (2018) but were not graphed. They were re-analyzed and their admixture components were illustrated in a bar graph in Wang et al (2018: figure 2c), but they are not the principal focus of any published study. All of the authors who examined them agreed that these three Khvalynsk individuals, dated about 4500 BC, showed EHG ancestry admixed substantially with CHG, and not a trace of Anatolian Farmer ancestry, so the CHG was a Hotu-Cave or Kotias-Cave type of un-admixed CHG. The proportion of CHG in the Wang et al. (2018) bar graphs is about 20-30% in two individuals, substantially less CHG than in Yamnaya; but the third Khvalynsk individual had more than 50% CHG, like Yamnaya. The ca. 30 additional unpublished individuals from three middle Volga Eneolithic cemeteries, including Khvalynsk, preliminarily show the same admixed EHG/CHG ancestry in varying proportions. Most of the males belonged to Ychromosome haplogroup R1b1a, like almost all Yamnaya males, but Khvalynsk also had some minority Y-chromosome haplogroups (R1a, Q1a, J, I2a2) that do not appear or appear only rarely (I2a2) in Yamnaya graves. The Journal of Indo-European Studies Archaeology, Genetics, and Language in the Steppes 13 Wang et al. (2018) discovered that this middle Volga mating network extended down to the North Caucasian steppes, where at cemeteries such as Progress-2 and Vonyuchka, dated 4300 BC, the same Khvalynsk-type ancestry appeared, an admixture of CHG and EHG with no Anatolian Farmer ancestry, with steppe-derived Y-chromosome haplogroup R1b. These three individuals in the North Caucasus steppes had higher proportions of CHG, overlapping Yamnaya. Without any doubt, a CHG population that was not admixed with Anatolian Farmers mated with EHG populations in the Volga steppes and in the North Caucasus steppes before 4500 BC. We can refer to this admixture as pre-Yamnaya, because it makes the best currently known genetic ancestor for EHG/CHG R1b Yamnaya genomes. The Progress-2 individuals from North Caucasus steppe graves lived not far from the pre-Maikop farmers of the Belaya valley, but they did not exchange mates, according to their DNA. The hunter-fisher camps that first appeared on the lower Volga around 6200 BC could represent the migration northward of un-admixed CHG hunter-fishers from the steppe parts of the southeastern Caucasus, a speculation that awaits confirmation from aDNA. After 5000 BC domesticated animals appeared in these same sites in the lower Volga, and in new ones, and in grave sacrifices at Khvalynsk and Ekaterinovka. CHG genes and domesticated animals flowed north up the Volga, and EHG genes flowed south into the North Caucasus steppes, and the two components became admixed. After approximately 4500 BC the Khvalynsk archaeological culture united the lower and middle Volga archaeological sites into one variable archaeological culture that kept domesticated sheep, goats, and cattle (and possibly horses). In my estimation, Khvalynsk might represent the oldest phase of PIE. Anatolian Farmer ancestry and Yamnaya origins The Eneolithic Volga-North Caucasus mating network (Khvalynsk/Progress-2 type) exhibited EHG/CHG admixtures and Y-chromosome haplogroups similar to Yamnaya, but without Yamnaya’s additional Anatolian Farmer ancestry. Were there other steppe populations that showed the same Volume 47, Number 1 & 2, Spring/Summer 2019 14 David W. Anthony EHG/CHG admixture? And when did Anatolian Farmer ancestry appear in the steppes? The Eneolithic Dnieper populations, concentrated around the Dnieper Rapids, were not good candidates for the ancestors of Yamnaya, because they lacked any CHG component. All of them were admixtures of EHG (primarily) and WHG (minor components). Among 32 published individuals from three Eneolithic cemeteries (Dereivka-1, Vil’nyaka, and Vovnigi) in the Dnieper River valley dated 5100-4400 BC, assigned to the Dnieper-Donets culture, there were no individuals with CHG ancestry. The steppe Y-chromosome haplogroup R1b1a was frequent at Dereivka-1, as in Yamnaya, but at the two other cemeteries on the Dnieper Rapids, Vil’nyanka and Vovigni, all the males were I2a2a. This was a common WHG haplogroup but only one Yamnaya male from Kalmykia has up to now been identified as I2a2. The Dnieper Rapids, flooded today by dams, were strategic fishing places and river crossings, and were the first place in the Pontic–Caspian steppes to attract populations that created formal cemeteries, beginning in the early Mesolithic about 11,000-10,500 BC (Lillie et al. 2009; Jones et al 2017: Supp. Mat. p.9). Like the Mesolithic and Neolithic populations here, the Eneolithic populations of Dnieper-Donets II type seem to have limited their mating network to the rich, strategic region they occupied, centered on the Rapids. The absence of CHG shows that they did not mate frequently if at all with the people of the Volga steppes, a surprising but undeniable discovery. Archaeologists have seen connections in ornament types and in some details of funeral ritual between Dnieper-Donets cemeteries of the Mariupol-Nikol’skoe type and cemeteries in the middle Volga steppes such as Khvalynsk and S’yez’zhe (Vasiliev 1981:122-123). Also their cranio-facial types were judged to be similar ((Bogdanov and Khokhlov 2012:212). So it it surprising that their aDNA does not indicate any genetic admixture with Khvalynsk or Progress-2. Also, neither they nor the Volga steppe Eneolithic populations showed any Anatolian Farmer ancestry. By 5000 BC permanent agricultural towns stood on tells throughout the lower Danube valley and Balkans, not far from the Pontic steppes. Large Cucuteni-Tripol’ye towns constructed The Journal of Indo-European Studies Archaeology, Genetics, and Language in the Steppes 15 of two-story buildings dotted the upland rainfall agriculture zone in the eastern Carpathian piedmont, overlooking the coastal Black Sea steppes. Yet among 48 individuals with whole-genome aDNA from 16 Neolithic and Copper Age cemeteries in Bulgaria and Romania dated 5800-4300 BC, only three showed any ancestry from a steppe mating network (Mathieson et al. 2018). Around 95% of the southeastern European farmer population tested had no steppe relatives over a period of 1500 years. They must have actively avoided marriage with steppe people, a rule broken only among the elite towards the end of the Eneolithic. All three of the steppe-admixed exceptions were from the Varna region (Mathieson et al. 2018). One of them was the famous “golden man’ at Varna (Krause et al. 2016), Grave 43, whose steppe ancestry was the most doubtful of the three. If he had steppe ancestry, it was sufficiently distant (five+ generations before him) that he was not a statistically significant outlier, but he was displaced in the steppe direction, away from the central values of the majority of typical Anatolian Farmers at Varna and elsewhere. The other two, at Varna (grave 158, a 5-7-year-old girl) and Smyadovo (grave 29, a male 20-25 years old), were statistically significant outliers who had recent steppe ancestry (consistent with grandparents or great-grandparents) of the EHG/CHG Khvalynsk/Progress-2 type, not of the Dnieper Rapids EHG/WHG type. Again, this is surprising, because the Volga is much farther away from Varna than the Dnieper. All three graves were unusually wellequipped with typical Varna pottery and ornaments, and grave 43 was spectacularly rich. Steppe people occasionally became the parents of children whose local parents belonged to Old European elite families, presumably as the result of arrangements tied to political and economic negotiations. But the children were kept in the tell towns and lived and died there. Aside from these three elite-looking Varna-region individuals dated 4650-4450 BC (Krause et al. 2016; Mathieson et al. 2018: Supplementary Materials), the majority of Eneolithic farmers who lived near the steppe region had no steppe relatives, mirroring the absence of Anatolian Farmer ancestry in Eneolithic steppe cemeteries. Volume 47, Number 1 & 2, Spring/Summer 2019 16 David W. Anthony This is surprising because many ornaments made of Balkan-derived copper reached the Volga, indicating some kind of exchange system (or looting?) with Balkan partners. At Khvalynsk, 373 copper ornaments were recovered from 27 graves among 199 excavated , the largest copper assemblage from a fifth-millennium BC site anywhere in the steppes (Agapov 2010). Most of it is argued to be Balkan in origin. Balkan copper also reached Svobodnoe, a site related to Meshoko (Courcier 2014: 586) and Progress-2-type steppe graves in the North Caucasus (Korenevskii 2016). Ornaments of Balkan copper, exotic ornamental shells (Glycemeris, Antalis), long lamellar flint blades, polished stone maces, and perhaps people and animals were exchanged across the Pontic–Caspian steppes from Varna to Khvalynsk and Svobodnoe between 4500-4200 BC. Steppe type graves appeared on the fringes of the lower Danube valley, at Suvorovo and Giurgiule
ti. Then about 4300-4200 BC the Varna/Karanovo VI-era tell towns of the lower Danube valley and the Balkans suffered a localized but sudden and total collapse resulting in the end of tell settlements and of most of their material culture traditions. The culture that replaced them in the lower Danube valley, Cernavoda I, was relatively impoverished and showed a mixture of steppe and Old European customs. I believe that the Suvorovo-Cernavoda I movement into the lower Danube valley and the Balkans about 4300 BC separated early PIE-speakers (pre-Anatolian) from the steppe population that stayed behind in the steppes and that later developed into late PIE and Yamnaya. This archaeological transition marked the breakdown of the mating barrier between steppe and Anatolian Farmer mating networks. After this 4300-4200 BC event, Anatolian Farmer ancestry began to pop up in the steppes. The currentlyoldest sample with Anatolian Farmer ancestry in the steppes in an individual at Aleksandriya, a Sredni Stog cemetery on the Donets in eastern Ukraine. Sredni Stog has often been discussed as a possible Yamnaya ancestor in Ukraine (Anthony 2007: 239254). The single published grave is dated about 4000 BC (4045– 3974 calBC/ 5215±20 BP/ PSUAMS-2832) and shows 20% Anatolian Farmer ancestry and 80% Khvalynsk-type steppe ancestry (CHG&EHG). His Y-chromosome haplogroup was R1a-Z93, similar to the later Sintashta culture and to South The Journal of Indo-European Studies Archaeology, Genetics, and Language in the Steppes 17 Asian Indo-Aryans, and he is the earliest known sample to show the genetic adaptation to lactase persistence (I3910
T). Another pre-Yamnaya grave with Anatolian Farmer ancestry was analyzed from the Dnieper valley at Dereivka, dated 3600-3400 BC (grave 73, 3634–3377 calBC/ 4725±25 BP/ UCIAMS-186349). She also had 20% Anatolian Farmer ancestry, but she showed less CHG than Aleksandriya and more Dereivka-1 ancestry, not surprising for a Dnieper valley sample, but also showing that the old fifth-millennium-type EHG/WHG Dnieper ancestry survived into the fourth millennium BC in the Dnieper valley (Mathieson et al. 2018). The 10-18% Anatolian Farmer ancestry that was widespread in Yamnaya was either a slight decline from or about the same as these older samples dated 4000-3500 BC in Ukraine. The west-to-east gene flow that began after the 43004200 BC collapse could have continued into the Yamnaya period. Part of the WHG that Wang et al. (2018) detected in Yamnaya genomes could have been picked up in the Dnieper valley, where many Dnieper-Donets individuals had WHG ancestry, possibly lessening the necessity for mate exchanges with Globular Amphorae. Probably, late PIE (Yamnaya) evolved in the same part of the steppes—the Volga-Caucasus steppes between the lower Don, the lower and middle Volga, and the North Caucasus piedmont—where early PIE evolved, and where appropriate EHG/CHG admixtures and Y-chromosome haplogroups were seen already in the Eneolithic (without Anatolian Farmer). There have always been archaeologists who argued for an origin of Yamnaya in the Volga steppes, including Gimbutas (1963), Merpert (1974), and recently Morgunova (2014), who argued that this was where Repin-type ceramics, an important early Yamnaya pottery type, first appeared in dated contexts before Yamnaya, about 3600 BC. The genetic evidence is consistent with Yamnaya EHG/CHG origins in the VolgaCaucasus steppes. Also, if contact with the Maikop culture was a fundamental cause of the innovations in transport and metallurgy that defined the Yamnaya culture, then the lower Don-North Caucasus-lower Volga steppes, closest to the North Caucasus, would be where the earliest phase is expected. Volume 47, Number 1 & 2, Spring/Summer 2019 18 David W. Anthony I would still guess that the Darkveti-Meshoko culture and its descendant Maikop culture established the linguistic ancestor of the Northwest Caucasian languages in approximately the region where they remained. I also accept the general consensus that the appearance of the hierarchical Maikop culture about 3600 BC had profound effects on preYamnaya and early Yamnaya steppe cultures. Yamnaya metallurgy borrowed from the Maikop culture two-sided molds, tanged daggers, cast shaft hole axes with a single blade, and arsenical copper. Wheeled vehicles might have entered the steppes through Maikop, revolutionizing steppe economies and making Yamnaya pastoral nomadism possible after 3300 BC. So it is still possible that steppe people interacted as raiders and traders and perhaps even political clients of the Maikop people, with interaction intense enough to make leading political figures in the pre-Yamnaya steppes bilingual in the Maikop (Northwest Caucasian?) language. Some Maikop women might also have become the wives of some preYamnaya men. If their speech was copied by others around them, the linguistic exchanges and interferences suggested by Bomhard could have occurred and spread without an equally large exchange of mates. But if the interpretations presented here are supported, mate exchanges between Maikop and preYamnaya or Yamnaya people were few in number, rare in frequency, and when they did happen, involved primarily Maikop women, not men. If more mating had occurred, we would see more EHG among the Maikop genomes and more Anatolian Farmer among Yamnaya steppe genomes than we do see. Of course another, final, possibility, consistent with the archaeological and genetic evidence presented here, is that there were two phases of interference from Caucasian languages in two periods. 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