Sea Peoples behind Philistines were Aegeans, including R1b-M269 lineages

New open access paper Ancient DNA sheds light on the genetic origins of early Iron Age Philistines, by Feldman et al. Science Advances (2019) 5(7):eaax0061.

Interesting excerpts (modified for clarity, emphasis mine):

Here, we report genome-wide data from human remains excavated at the ancient seaport of Ashkelon, forming a genetic time series encompassing the Bronze to Iron Age transition. We find that all three Ashkelon populations derive most of their ancestry from the local Levantine gene pool. The early Iron Age population was distinct in its high genetic affinity to European-derived populations and in the high variation of that affinity, suggesting that a gene flow from a European-related gene pool entered Ashkelon either at the end of the Bronze Age or at the beginning of the Iron Age. Of the available contemporaneous populations, we model the southern European gene pool as the best proxy for this incoming gene flow. Last, we observe that the excess European affinity of the early Iron Age individuals does not persist in the later Iron Age population, suggesting that it had a limited genetic impact on the long-term population structure of the people in Ashkelon.

philistines-pca
Ancient genomes (marked with color-filled symbols) projected onto the principal components inferred from present-day west Eurasians (gray circles). The newly reported Ashkelon populations are annotated in the upper corner.

Genetic discontinuity between the Bronze Age and the early Iron Age people of Ashkelon

In comparison to ASH_LBA, the four ASH_IA1 individuals from the following Iron Age I period are, on average, shifted along PC1 toward the European cline and are more spread out along PC1, overlapping with ASH_LBA on one extreme and with the Greek Late Bronze Age “S_Greece_LBA” on the other. Similarly, genetic clustering assigns ASH_IA1 with an average of 14% contribution from a cluster maximized in the Mesolithic European hunter-gatherers labeled “WHG” (shown in blue in Fig. 2B) (15, 22, 26). This component is inferred only in small proportions in earlier Bronze Age Levantine populations (2 to 9%).

In agreement with the PCA and ADMIXTURE results, only European hunter-gatherers (including WHG) and populations sharing a history of genetic admixture with European hunter-gatherers (e.g., as European Neolithic and post-Neolithic populations) produced significantly positive f4-statistics (Z ≥ 3), suggesting that, compared to ASH_LBA, ASH_IA1 has additional European-related ancestry.

We find that the PC1 coordinates positively correlate with the proportion of WHG ancestry modeled in the Ashkelon individuals, suggesting that WHG reasonably tag a European-related ancestral component within the ASH_IA1 individuals.

philistines-admixture
We plot the ancestral proportions of the Ashkelon individuals inferred by qpAdm using Iran_ChL, Levant_ChL, and WHG as sources ±1 SEs. P values are annotated under each model. In cases when the three-way model failed (χ2P < 0.05), we plot the fitting two-way model. The WHG ancestry is necessary only in ASH_IA1.

The best supported one (χ2P = 0.675) infers that ASH_IA1 derives around 43% of ancestry from the Greek Bronze Age “Crete_Odigitria_BA” (43.1 ± 19.2%) and the rest from the ASH_LBA population.

(…) only the models including “Sardinian,” “Crete_Odigitria_BA,” or “Iberia_BA” as the candidate population provided a good fit (χ2P = 0.715, 49.3 ± 8.5%; χ2P = 0.972, 38.0 ± 22.0%; and χ2P = 0.964, 25.8 ± 9.3%, respectively). We note that, because of geographical and temporal sampling gaps, populations that potentially contributed the “European-related” admixture in ASH_IA1 could be missing from the dataset.

The transient impact of the “European-related” gene flow on the Ashkelon gene pool

The ASH_IA2 individuals are intermediate along PC1 between the ASH_LBA ones and the earlier Bronze Age Levantines (Jordan_EBA/Lebanon_MBA) in the west Eurasian PCA (Fig. 2A). Notably, despite being chronologically closer to ASH_IA1, the ASH_IA2 individuals position closer, on average, to the earlier Bronze Age individuals.

philistines-y-dna
See more information on Y-DNA SNP calls, including ASH067 as R1b-M269 (xL151).

The transient excess of European-related genetic affinity in ASH_IA1 can be explained by two scenarios. The early Iron Age European-related genetic component could have been diluted by either the local Ashkelon population to the undetectable level at the time of the later Iron Age individuals or by a gene flow from a population outside of Ashkelon introduced during the final stages of the early Iron Age or the beginning of the later Iron Age.

By modeling ASH_IA2 as a mixture of ASH_IA1 and earlier Bronze Age Levantines/Late Period Egyptian, we infer a range of 7 to 38% of contribution from ASH_IA1, although no contribution cannot be rejected because of the limited resolution to differentiate between Bronze Age and early Iron Age ancestries in this model.

Hg. R1b-M269 and the Aegean

I already predicted this relationship of Philistines and Aegeans (Greeks in particular) months ago, based on linguistics, archaeology, and phylogeography, although it was (and still is) yet unclear if these paternal lineages might have come from other nearby populations which might be descended from Common Anatolians instead, given the known intense contacts between Helladic and West Anatolian groups.

luwian-civilization-sea-peoples
The alternative view: The Sea Peoples can be traced back to the Aegean, so they could also have consisted of Luwian petty kingdoms, who had formed an alliance and attacked Hatti from the south.

The deduction process for the Greek connection was quite simple:

Palaeo-Balkan populations

We know that R1b-Z2103 expanded with Yamna, including West Yamna settlers: they appear in Vučedol, which means they formed part of the earliest expansion waves of Yamna settlers into the Carpathian Basin, and they also appear scattered among Bell Beakers (apart from dominating East Yamna and Afanasevo), which suggests that they were possibly one of the most successful lineages during the late Repin/early Yamna expansion.

The “Steppe ancestry” associated with I2a-L699 samples among Balkan BA peoples may have also been associated with recent Bronze Age expansions, and this haplogroup’s presence among modern Balkan peoples may also suggest that it expanded with Palaeo-Balkan languages. Nevertheless, we don’t know which specific lineages and “Steppe ancestry” they represent, sadly.

These samples may well be related to remnants of previous Balkan populations like Cernavodă or Ezero, because there has been no peer-reviewed attempt at distinguishing Khvalynsk-/Novodanilovka- from Sredni Stog- from Yamnaya-related populations (see here), and some groups that are associated with this ancestry, like Corded Ware, are known to be culturally distinct from Yamna.

In any case, Proto-Greeks from the southern Balkans (say, Sitagroi IV and related groups) are probably going to show, based on Palaeo-Balkan substrate and Pre-Greek substrate and on the available Mycenaean samples, a process of decreasing proportion of R1b-Z2103 lineages relative to local ones, and a relatively similar cline of Yamna:EEF ancestry from northern to southern areas, at least in the periods closest to the Yamna expansion.

NOTE. The finding of “archaic” R1b-L389 (R1b-V1636) and R1a-M198 subclades among modern Greeks and the likely Neolithic origin of these paternal lineages around the Caucasus suggest that their presence in Greece may be from any of the more recent migrations that have happened between Anatolia and the Balkans, especially during the Common Era, rather than Indo-Anatolian migrations; probably very very recently.

-chalcolithic-late-balkans
Bronze Age cultures in the Balkans and the Aegean. See full map including ancient samples with Y-DNA, mtDNA, and ADMIXTURE.

Minoans and haplogroup J

In the Aegean, it is already evident that the population changed language partly through cultural diffusion, probably through elite domination of Proto-Greek speakers. Whether that happened before the invasion into the Greek Peninsula or after it is unclear, as we discussed recently, because we only have one reported Y-chromosome haplogroup among Mycenaeans, and it is J (probably continuing earlier lineages).

Now we have more samples from the so-called Emporion 2 cluster in Olalde et al. (2019), which shows Mycenaean-like eastern Mediterranean ancestry and 3 (out of 3) samples of haplogroup J, which – given the origin of the colony in Phocea – may be interpreted as the prevalence of West Anatolian-like ancestry and lineages in the eastern part of the Aegean (and possibly thus south Peloponnese), in line with the modern situation.

NOTE. It does not seem likely that those R or R1b-L23 samples from the Emporion 1 cluster are R1b-Z2103, based on their West European-like ancestry, although they still may be, because – as we know – ancestry (unlike haplogroup) changes too easily to interpret it as an ancestral ethnolinguistic marker.

anatolia-greek-aegean
PCA of ancient samples related to the Aegean, with Minoans, Mycenaeans (including the Emporion 2 cluster in the background) Anatolia N-Ch.-BA and Levantine BA-LBA populations, including Tel Shadud samples. See more PCAs of ancient Eurasian populations.

Greeks and haplogroup R1b-M269

Therefore, while the presence of R1b-Z2103 among ancient Balkan peoples connected to the Yamna expansion is clear, one might ask if R1b-Z2103 really spread up to the Peloponnese by the time of the Mycenaean Civilization. That has only one indirect answer, and it’s most likely yes.

We already had some R1b-Z2103 among Thracians and around the Armenoid homeland, which offers another clue at the migration of these lineages from the Balkans. The distribution of different “archaic” R1b-Z2103 subclades among modern Balkan populations and around the Aegean offered more support to this conclusion.

But now we have two interesting ancient populations that bear witness to the likely intrusion of R1b-M269 with Proto-Greeks:

An Ancient Greek of hg. R1b

A single ancient sample supports the increase in R1b-Z2103 among Greeks during the “Dorian” invasions that triggered the Dark Ages and the phenomenon of the Aegean Sea Peoples. It comes from a Greek lab study, showing R1b1b (i.e. R1b-P297 in the old nomenclature) as the only Y-chromosome haplogroup obtained from the sampling of the Gulf of Amurakia ca. 470-30 BC, i.e. before the Roman foundation of Nikopolis, hence from people likely from Anaktorion in Ancient Acarnania, of Corinthian origin.

ancient-greeks-y-dna-mtdna

Even with the few data available – and with the caution necessary for this kind of studies from non-established labs, which may be subject to many different kinds of errors – one could argue that the western Greek areas, which received different waves of migrants from the north and shows a higher distribution of R1b-Z2103 in modern times, was probably more heavily admixed with R1b-Z2103 than southern and eastern areas, which were always dominated by Greek-speaking populations more heavily admixed with locals.

The Dorian invasion and the Greek Dark Ages may thus account for a renewed influx of R1b-Z2103 lineages accompanying the dialects that would eventually help form the Hellenic Koiné. In a sense, it is only natural that demographically stronger populations around the Bronze Age Aegean would suffer a limited (male) population replacement with the succeeding invasions, starting with a higher genetic impact in the north-west and diminishing as they progressed to the south and the east, coupled with stepped admixture events with local populations.

This would be therefore the late equivalent of what happened at the end of the 3rd millennium BC, with Mycenaeans and their genetic continuity with Minoans.

pre-greek-ssos
Distribution of Pre-Greek place-names ending in -ssos/-ssa or -sos/-sa. See original images and more on the south/east cline distribution of Pre-Greek place-names here.

Sea peoples of hg. R1b-M269

Thanks to Wang et al. (2018) supplementary materials we knew that one of the two Levantine LBA II samples from Tel Shadud (final 13th–early 11th c. BC) published in van den Brink (2017) was of hg. R1b-M269 – in fact, the one interpreted as a Canaanite official residing at this site and emulating selected funerary aspects of Egyptian mortuary culture.

Both analyzed samples, this elite individual and a commoner of hg. J buried nearby, were genetically similar and indistinguishable from local populations, though:

Principal Components Analysis of L112 and L126 was carried out within the framework described in Lazaridis et al. (2016). This analysis showed that the two individuals cluster genetically, with similar estimated proportions of ancestry from diverse West Eurasian ancestral sources. These results are consistent with the hypothesis that they derive from the same population, or alternatively that they derive from two quite closely related populations.

We know that ancestry changes easily within a few generations, so there was not much information to go on, except for the fact that – being R1b-M269 – this individual could trace his paternal ancestor at some point to Proto-Indo-Europeans.

One might think that, because many haplogroups in this spreadsheet were wrong, this is also wrong; nevertheless, many haplogroups are correctly identified by Yleaf, and finding R1b-M269 in the Levant after the expansion of Sea Peoples could not be that surprising, because they were most likely related to populations of the Aegean Sea. Any other related hg. R1b (R1b-M73, R1b-V88, even R1b-V1636) wouldn’t fit as well as R1b-M269.

sea-peoples-egypt-rameses-iii

However, the early expansion of Proto-Indo-Aryans into the Middle East, as well as the later expansion of Armenians from the Balkans through Anatolia and of West Iranians from the east may have all potentially been related to this sample. But still, the previous linguistic and archaeological theories concerning the Philistines and the expansion of Sea Peoples in the Levant made this sample a likely (originally) Greek “Dorian” lineage, rather than the other (increasingly speculative) alternatives.

In any case, it was obvious to anyone – that is, to anyone with a minimum knowledge of how population genomics works – that just the two samples from van den Brink (2017) couldn’t be used to get to any conclusions about the ancestral origin of these individuals (or their differences) beyond Levantine peoples, because their ancestry was essentially (i.e. statistically) the same as the other few available ancient samples from nearby regions and similar periods.

If anything, the PCA suggested an origin of the R1b sample closer to Aegean populations relative to the J individual (see PCA above), and this should have been supported also by amateur models, without any possible confirmation (as with the ASH_IA2 cluster in this paper). However, if you have followed online discussions of Tel Shadud R1b-M269 sample since it was mentioned first on Eupedia months ago – including another wave of misguided speculation based on the ancestry of both individuals triggered by a discussion on this blog -, you have once more proof of how misleading ancestry analyses can be in the wrong hands.

NOTE. This is the Nth proof (and that only in 2019) of how it’s best to just avoid amateur analyses and interpretations altogether, as I did in the recent publication of the books. All those who didn’t take into account whatever was commented about the ancestry of these samples haven’t lost a single bit of relevant information on Levantine peoples, and have had more time for useful reads, compared to those dedicated to endless void speculation, once again gone awfully wrong, as does everything related to cocky ancient DNA crackpottery 😉

bronze-age-late-aegean
Late Bronze Age population movements in the Eastern Mediterranean and the Middle East. See full map including ancient DNA samples with Y-DNA, mtDNA, and ADMIXTURE.

Admittedly, though, even accepting the evident Mediterranean origin of this lineage, one could have argued that this sample may have been of R1b-L151 subclade, if one were inclined to support the theory that Italic peoples were behind Sea Peoples expanding east – and consequently that the ancestors of Etruscans had migrated eastward into the Aegean (e.g. into Lemnos), so that it could be asserted that Tyrsenian might have been a remnant language of an ancient population of northern Italy.

Philistines

Fortunately, some of the samples recovered in Feldman et al. (2019) that could be analyzed (those of the cluster ASH_IA1) offer a very specific time frame where European ancestry appeared (ca. 1250 BC) before it subsequently became fully diluted (as seen in cluster ASH_IA2) among the prevalent Levantine ancestry of the area.

Also fortunately, this precise cluster shows another R1b-M269 sample, likely R1b-Z2103 (because it is probably xL151), and this sample together with others from the same cluster prove that the ancestry related to the original southern European incomers was:

  1. Recent, related thus to LBA population movements, as expected; and
  2. More closely related to coeval Aegeans, including Mycenaeans with Steppe-related ancestry.

NOTE. I say “fortunately” because, as you can imagine if you have dealt with amateurish discussions long enough, without this cluster with evident Aegean ancestry and the R1b-M269 (Z2103) sample precisely associated to it, some would enter again in endless comment loops created by ancestry magicians, showing how Aegean peoples were not behind Sea Peoples, or not behind Philistines, or not behind the R1b-M269 among Philistines, depending on their specific agendas.

aegean-sea-peoples
Map of the Sea People invasions in the Aegean Sea and Eastern Mediterranean at the end of the Late Bronze Age (blue arrows).. Some of the major cities impacted by the raids are denoted with historical dates. Inland invasions are represented by purple arrows. From Kaniewski et al. (2011). Some of the major cities impacted by the raids are denoted with historical dates. Inland invasions are represented by purple arrows.

The results of the paper don’t solve the question of the exact origin of all Sea Peoples (not even that of Philistines), but it is quite clear that most of those forming this seafaring confederation must have come from sites around the Aegean Sea. This supports thus the traditional origin attributed to them, including a hint at the likely expansion of Eastern Mediterranean ancestry and lineages into the Italian Peninsula precisely from the Aegean, as some oral communications have already disclosed.

As an indirect conclusion from the findings in this paper, then, we can now more confidently support that Tyrsenian speakers most likely expanded into the Appenines and the Alps originally from a Tyrsenian-speaking LBA population from Lemnos, due to the social unrest in the whole Aegean region, and might have become heavily admixed with local Italic peoples quite quickly, as it happened with Philistines, resulting in yet another case of language expansion through (the simplistically called) elite domination.

Conclusion

Even more interesting than these specific findings, this paper confirms yet another hypothesis based on phylogeography, and proves once again two important starting points for ancient DNA interpretation that I have discussed extensively in this blog:

  • The rare R1b-M269 Y-chromosome lineage of Tel Shadud offered ipso facto the most relevant clue about the ancestral geographical origin of this Canaanite elite male’s paternal family, most likely from the north-west based on ancient phylogeography, which indirectly – in combination with linguistics and archaeology – supported the ancestral ethnolinguistic identification of Philistines with the Aegean and thus with (a population closest to) Ancient Greeks.
  • Ancestry analyses are often fully unreliable when assessing population movements, especially when few samples from incomplete temporal-geographical transects are assessed in isolation, because – unlike paternal (and maternal) haplogroups – ancestry might change fully within a few generations, depending on the particular anthropological setting. Their investigation is thus bound by many limitations – of design, statistical, and anthropological (i.e. archaeological and linguistic) – which are quite often not taken into account.

These cornerstones of ancient DNA interpretation have been already demonstrated to be valid not only for Levantine populations, as in this case, but also for Balkan peoples, for Bell Beakers, for steppe populations (like Khvalynsk, Sredni Stog, Yamna, Corded Ware), for Basques, for Balto-Slavs, for Ugrians and Samoyeds, and for many other prehistoric peoples.

I rest my case.

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).

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Pasture usage by ancient pastoralists in Middle and Late Bronze Age Kazakhstan

bestamak-lisakovsk-study-area

Open access Pasture usage by ancient pastoralists in the northern Kazakh steppe informed by carbon and nitrogen isoscapes of contemporary floral biomes, by Miller et al. Archaeol Anthropol Sci (2018).

Interesting excerpts (emphasis mine):

Bronze age settlement, society, and subsistence in the northern Kazakh steppe

The Middle to Late Bronze Age (2200 to 1400 cal BCE) in the northern Kazakh steppe encompassed a major shift in settlement patterns from semi-sedentary pastoralism to more dispersed, mobile lifeways engaged in pastoral nomadism (Tkacheva 1999; Grigory’ev 2002; Koryakova and Epimakhov 2007; Kuz’mina 2007; Tkacheva and Tkachev 2008). Middle Bronze Age (2200 to 1700 cal BCE) settlements had large enclosures consisting of an earthen wall and ditch. Inside the enclosure, earthen domestic structures with shared walls (numbering from 30 to 60) housed an estimated 200 to 700 individuals (Gening et al. 1992; Grigor’yev 2002; Anthony 2007; Kohl 2007; Koryakova and Epimakhov 2007; Hanks 2009; Batanina and Hanks 2013). MBA settlements were repeatedly occupied, evidenced by successive building phases that added structures and enlarged enclosures. Aggregated MBA sites are situated between 40 and 60 km apart, and landscapes between enclosed settlements may have been territories of particular settlements (Epimakhov 2002; Zdanovich and Batanina 2002; Merrony et al. 2009; Stobbe et al. 2016). While there is currently no archeological evidence for structures such as animal corrals or walls outside of MBA settlement enclosures, open areas within settlements may have been used to house livestock. Reconstructions of landscape use in the vicinity of MBA sites determined that pastures within 4 km of the site could have supported herd sizes large enough to sustain sedentary livestock herders (Stobbe et al. 2016). During the subsequent Late Bronze Age (1800 to 1400 cal BCE) settlements were more dispersed across the landscape and significantly smaller, consisting of fewer than 20 dwellings, further lacking enclosures and building phases (Kuz’mina 2007:36–8; Zakh and Ilyushina 2010). This shift in settlement size and distribution has been interpreted to indicate the emergence of nomadic pastoralism and the intensification of long-distance mobility (Tkacheva 1999; Grigory’ev 2000; Kuz’mina 2007; Tkacheva and Tkachev 2008).

MBA communities engaged in pastoralism and supplemented their diets with wild plants and wild game (Krause and Koryakova 2013; Ventresca Miller et al. 2014a; Hanks et al. 2018). A variety of wild plants have been recovered during flotation, but so far, domesticated grains have not been recovered (Krause and Koryakova 2013; Ng 2013; Hanks et al. 2018). Carbon and nitrogen stable isotope analyses of bone collagen indicate that human dietary intake in the MBA focused on terrestrial animal protein, likely in the form of meat and milk, which was supplemented by locally available fish and wild plants (Ventresca Miller et al. 2014a; Hanks et al. 2018). While the subsequent LBA has been interpreted as a shift to nomadic pastoralism, little data is available regarding landscape use or herd management strategies for this period. Paleodietary studies suggest that human diets during the LBA focused on pastoral products and were supplemented by wild plants, fish, and wild animals (Ventresca Miller et al. 2014a, 2014b).

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Location of the archeological sites of Bestamak (MBA), Kamennyi Ambar (MBA), Bolshekaragansky (MBA), and Lisakovsk (LBA)

Conclusions

A major shift in patterns of settlement occurred at the Middle to Late Bronze Age transition, from large semi-sedentary populations in enclosed settlements to smaller populations in open settlements dispersed across the landscape. Scholars have suggested that animal management strategies also changed at this time from semi-sedentary pastoralism to more mobile forms of pastoral nomadism. However, our findings suggest that livestock management practices did not shift in concert with social landscapes, demonstrating consistency in pastoral adaptations through time in the region. Similar isotopic patterning between livestock during the MBA and LBA across several sites in the CES indicates that there were no changes across time in pasture usage patterns. Among ancient livestock, differences in δ13C and δ15N values between horses and ruminants (cattle, sheep, goat) strongly suggest that livestock were grazed pastures either extensively or intensively, respectively. Horses grazed in open steppe areas or intermittently in areas with well-watered soils that lacked salinity, likely staying well outside of settlements. In contrast, cattle and sheep/goat grazed in pastures across multiple zones, both near the settlement and in non-local pastures that were grazed intensively. A wider range of δ13C and δ15N values among ruminants at Kamennyi Ambar (MBA) suggests that aggregated human populations may have had larger herds, some of which accessed non-local pastures outside of the easily accessible territories surrounding enclosed sites. Continued research on the isotopic composition of vegetation surrounding Bronze Age sites should clarify patterns of landscape use between MBA sites.

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