Yersinia pestis, the etiologic agent of plague, is a bacterium associated with wild rodents and their fleas. Historically it was responsible for three pandemics: the Plague of Justinian in the 6th century AD, which persisted until the 8th century [ 1 ]; the renowned Black Death of the 14th century [ 2, 3 ], with recurrent outbreaks until the 18th century [ 4 ]; and the most recent 19th century pandemic, in which Y. pestis spread worldwide [ 5 ] and became endemic in several regions [ 6 ]. The discovery of molecular signatures of Y. pestis in prehistoric Eurasian individuals and two genomes from Southern Siberia suggest that Y. pestis caused some form of disease in humans prior to the first historically documented pandemic [ 7 ]. Here, we present six new European Y. pestis genomes spanning the Late Neolithic to the Bronze Age (LNBA; 4,800 to 3,700 calibrated years before present). This time period is characterized by major transformative cultural and social changes that led to cross-European networks of contact and exchange [ 8, 9 ]. We show that all known LNBA strains form a single putatively extinct clade in the Y. pestis phylogeny. Interpreting our data within the context of recent ancient human genomic evidence that suggests an increase in human mobility during the LNBA, we propose a possible scenario for the early spread of Y. pestis: the pathogen may have entered Europe from Central Eurasia following an expansion of people from the steppe, persisted within Europe until the mid-Bronze Age, and moved back toward Central Eurasia in parallel with human populations.
It seems that, notwithstanding the simplistic (white) arrows of steppe ancestry expansion shown in their map (see below), the actual expansion of Yersinia pestis might have in fact accompanied Yamna migrants from the Pontic-Caspian steppe into Early Bronze Age cultures from the Balkans, including Bell Beaker migrants, as the phylogenetic analysis and dates suggest – and as the potential arrows of the plague expansion in the map (in green) show.
Instead of warring nature, close ties, and mobility of Corded Ware peoples (reasons I used to justify the rapid spread of the disease among CWC groups), I guess it was rather the higher population density of SE Europecompared to the regions north of the loess belt, as well as the greater admixture of Yamna migrants with native SE European populations, the factors which might have helped expand the disease.
Nevertheless, lacking more data, it is unclear if the disease expanded with both steppe groups.
I already wrote about the concept of outlier in Human Ancestry, so I am not going to repeat myself. This is just an update of “outliers” in recent studies, and their potential origins (here I will repeat some of the examples):
Early Khvalynsk: the three samples from the Samara region have quite different positions in PCA, from nearest to EHG (of Y-DNA haplogroup R1a) to nearest to ANE ancestry (of Y-DNA haplogroup Q). This could represent the initial consequences of the second wave of ANE ancestry – as found later in Yamna samples from a neighbouring region -, possibly brought then by Eurasian migrants related to haplogroup Q.
With only 3 samples, this is obviously just a tentative explanation of the finds. The samples can only be reasonably said to show an unstable time for the region in terms of admixture (i.e. probably migration), judging by the data on PCA.
Ukraine Eneolithic samples offer a curious example of how the concept of outlier can change radically: from the third version (May 30th) of the preprint paper of Mathieson et al. (2017), when the Ukraine Eneolithic sample with steppe ancestry (and clustering with central European samples) was the ‘outlier’, to the fourth version (September 19th), when two samples with steppe ancestry clustering close to Corded Ware samples were now the ‘normal’ ones (i.e. those representing Ukraine Eneolithic population), and the outlier was the one clustering closely with Ukraine Mesolithic samples…
This is one of the funny consequences of the wrong interpretation of the ‘yamnaya component’, that made geneticists believe at first that, out of two samples (!), the ‘outlier’ was the one with ‘yamnaya’ ancestry, because this component would have been brought by an eastern immigrant from early Khvalynsk…
West Yamna (to insist on the same question, the ‘yamnaya’ component): we have only four western Yamna samples, two of them showing Anatolian Neolithic ancestry (one of them, from Ukraine, with a strong ‘southern’ drift). On the other hand, Corded Ware migrants do not show this. So we could infer that their migrations were not coetaneous: whereas peoples of Corded Ware culture expanded ca. 3300 BC to the north – in the natural corridor to the Baltic that has been proposed for this culture in Archaeology for decades (and that is well represented by Ukraine Eneolithic samples) -, peoples of Yamna culture expanded to the west, replacing the Ukraine Eneolithic population (i.e. probably those of ‘Proto-Corded Ware culture’), and eventually mixing with Balkan populations of Anatolian Neolithic ancestry.
Potapovka, Andronovo, and Srubna: while Potapovka clusters closely to the steppe, and Andronovo (like Sintashta) clusters closely to Corded Ware (i.e. Ukraine Neolithic / Central-East European), both have certain ‘outliers’ in PCA: the former has one individual clustering closely to Corded Ware, and the latter to the steppe. Both ‘outliers’ fit well with the interpretation of the recent mixture of Corded Ware peoples with steppe populations, and they offer a different image for the evolution of populations of Potapovka and Sintashta-Petrovka, potentially influencing their language. The position of Srubna samples, nearer to Sintashta and Andronovo (but occupying the same territory as the previous Potapovka) offers the image of a late westward conquest from Corded Ware-related populations.
Iron Age Bulgaria: a sample of haplogroup R1a-z93, with more ‘yamnaya’ ancestry than any other previous sample from the Balkans. For some, it might mean continuity from an older time. However – as with the Corded Ware outlier from Esperstedt before it – it is more likely a recent migrant from the steppe. The most likely origin of this individual is therefore people from the steppe, i.e. either the Srubna culture or a related group. Its relatively close cluster in PCA to certain recent Slavic populations can be interpreted in light of the multiple back and forth migrations in the region: of steppe populations to the west (Srubna, Cimmerians, Scythians, Sarmatians,…), and of Slavic-speaking populations:
And then rapidly expanding as a Proto-Slavic-speaking community from the steppe to the west.
Well-defined outliers are, therefore, essential to understand a recent history of admixture. On the other hand, the very concept of “outlier” can be a dangerous tool – when the lack of enough samples makes their classification as as such unjustified -, leading to the wrong interpretations.
It is unclear whether Indo-European languages in Europe spread from the Pontic steppes in the late Neolithic, or from Anatolia in the Early Neolithic. Under the former hypothesis, people of the Globular Amphorae culture (GAC) would be descended from Eastern ancestors, likely representing the Yamnaya culture. However, nuclear (six individuals typed for 597 573 SNPs) and mitochondrial (11 complete sequences) DNA from the GAC appear closer to those of earlier Neolithic groups than to the DNA of all other populations related to the Pontic steppe migration. Explicit comparisons of alternative demographic models via approximate Bayesian computation confirmed this pattern. These results are not in contrast to Late Neolithic gene flow from the Pontic steppes into Central Europe. However, they add nuance to this model, showing that the eastern affinities of the GAC in the archaeological record reflect cultural influences from other groups from the East, rather than the movement of people.
Excerpt, from the discussion:
In its classical formulation, the Kurgan hypothesis, i.e. a late Neolithic spread of proto-Indo-European languages from the Pontic steppes, regards the GAC people as largely descended from Late Neolithic ancestors from the East, most likely representing the Yamna culture; these populations then continued their Westward movement, giving rise to the later Corded Ware and Bell Beaker cultures. Gimbutas  suggested that the spread of Indo-European languages involved conflict, with eastern populations spreading their languages and customs to previously established European groups, which implies some degree of demographic change in the areas affected by the process. The genomic variation observed in GAC individuals from Kierzkowo, Poland, does not seem to agree with this view. Indeed, at the nuclear level, the GAC people show minor genetic affinities with the other populations related with the Kurgan Hypothesis, including the Yamna. On the contrary, they are similar to Early-Middle Neolithic populations, even geographically distant ones, from Iberia or Sweden. As already found for other Late Neolithic populations , in the GAC people’s genome there is a component related to those of much earlier hunting-gathering communities, probably a sign of admixture with them. At the nuclear level, there is a recognizable genealogical continuity from Yamna to Corded Ware. However, the view that the GAC people represented an intermediate phase in this large-scale migration finds no support in bi-dimensional representations of genome diversity (PCA and MDS), ADMIXTURE graphs, or in the set of estimated f3-statistics.
Together with Globular Amphora culture samples from Mathieson et al. (2017), this suggests that Kristiansen’s Indo-European Corded Ware Theory is wrong, even in its latest revised models of 2017.
On the other hand, the article’s genetic finds have some interesting connections in terms of mtDNA phylogeography, but without a proper archaeological model it is difficult to explain them.
Background: The Eneolithic (~5,500 yrBP) site of Verteba Cave in Western Ukraine contains the largest collection of human skeletal remains associated with the archaeological Cucuteni-Tripolye Culture. Their subsistence economy is based largely on agro-pastoralism and had some of the largest and most dense settlement sites during the Middle Neolithic in all of Europe. To help understand the evolutionary history of the Tripolye people, we performed mtDNA analyses on ancient human remains excavated from several chambers within the cave.
Results: Burials at Verteba Cave are largely commingled and secondary in nature. A total of 68 individual bone specimens were analyzed. Most of these specimens were found in association with well-defined Tripolye artifacts. We determined 28 mtDNA D-Loop (368 bp) sequences and defined 8 sequence types, belonging to haplogroups H, HV, W, K, and T. These results do not suggest continuity with local pre-Eneolithic peoples, but rather complete population replacement. We constructed maximum parsimonious networks from the data and generated population genetic statistics. Nucleotide diversity (π) is low among all sequence types and our network analysis indicates highly similar mtDNA sequence types for samples in chamber G3. Using different sample sizes due to the uncertainly in number of individuals (11, 28, or 15), we found Tajima’s D statistic to vary. When all sequence types are included (11 or 28), we do not find a trend for demographic expansion (negative but not significantly different from zero); however, when only samples from Site 7 (peak occupation) are included, we find a significantly negative value, indicative of demographic expansion.
Conclusions: Our results suggest individuals buried at Verteba Cave had overall low mtDNA diversity, most likely due to increased conflict among sedentary farmers and nomadic pastoralists to the East and North. Early Farmers tend to show demographic expansion. We find different signatures of demographic expansion for the Tripolye people that may be caused by existing population structure or the spatiotemporal nature of ancient data. Regardless, peoples of the Tripolye Culture are more closely related to early European farmers and lack genetic continuity with Mesolithic hunter-gatherers or pre-Eneolithic groups in Ukraine.
Genetic finds keep supporting the long-lasting cultural and linguistic frontier that Anthony (2007) – among others – asserted existed in the North-West Pontic steppe in the Mesolithic and Neolithic, between western steppe cultures and farmers, while it disproves Kristiansen’s theories of Sredni Stog expansion in Kurgan waves with a mixture of GAC and Trypillia within the Corded Ware culture:
Previous ancient DNA studies showed that hunter-gatherers before 6,500 yrBP in Europe commonly had haplogroups U, U4, U5, and H, whereas hunter-gatherers after 6,500 yrBP in Europe had less frequency of haplogroup H than before. Haplogroups T and K appeared in hunter-gatherers only after 6,500 yrBP, indicating a degree of admixture in some places between farmers and hunter-gatherers. Farmers before and after 6,500 yrBP in Europe had haplogroups W, HV*, H, T, K, and these are also found in individuals buried at Verteba Cave. Therefore, our data point to a common ancestry with early European farmers. Our data also suggest population replacement. Mathieson et al. analyzed a number of Neolithic Ukrainian samples (petrous bone) from several sites in southern, northern, and western Ukraine, dating to ~8,500 – 6,000 yrBP, and found exclusively U (U4 and U5) mtDNA lineages. It should be noted that ‘Neolithic’ in this context does not mean the adoption of agriculture, but rather simply coinciding with a change in material culture. They also analyzed several Trypillian individuals from Verteba Cave (different samples from the those included in this study). Similar to our findings, they found a wider diversity of mtDNA lineages, including H, HV, and T2b. These data, combined with our results, appear to confirm almost complete population replacement by individuals associated with the Tripolye Culture during the Middle to Late Neolithic.
The findings also hint to potential contacts of Yamna with Usatovo as predicted by Anthony (2007), or alternatively (lacking precise dates) to contacts with Corded Ware migrants:
Trypillians were very much a distinct people who most likely displaced 1 local hunter-gatherers with little admixture. Haplogroup W was also observed in several specimens deriving from Site G3. Although we are unsure if all of these haplogroups come from a single or multiple individuals, this observation is interesting in that it is relatively rare and isolated among Neolithic samples. It has, however, been found in samples dating to the Bronze Age. In the study by Wilde et al. , they found haplogroup W present in two samples from the Early Bronze Age associated with the Yamnaya and Usatovo cultures. The Usatovo culture (~ 3500 – 2500 BC) was found in Romania, Moldova, and southern Ukraine. It was the conglomeration of Tripolye and North Pontic steppe cultures. Therefore, this individual could link the Trypillian peoples to the Usatovo peoples and perhaps to the greater Yamnaya steppe migrations during the Bronze Age that lead to the Corded Ware Culture.
On the other hand, an article written in terms of mtDNA haplogroup frequencies seems to offer too little proof of anything today. The lack of Y-DNA haplogroups and data on admixture makes their interpretations provisional, subject to change when these further data are published. Also, radiocarbon dating is only confident for individuals of one site (site 7), dated ca. 5,500 cal BP, while “other chambers in the cave are not as confidently dated”…
Many researchers have pointed to the huge “megasites” and construction of fortifications as evidence of intergroup hostilities among the Late Neolithic Tripolye archaeological culture. However, to date, very few skeletal remains have been analyzed for the types of traumatic injury that serve as direct evidence for violent conflict. In this study, we examine trauma on human remains from the Tripolye site of Verteba Cave in western Ukraine. The remains of 36 individuals, including 25 crania, were buried in the gypsum cave as secondary interments. The frequency of cranial trauma is 30-44% among the 25 crania, six males, four females and one adult of indeterminate sex displayed cranial trauma. Of the 18 total fractures, 10 were significantly large and penetrating suggesting lethal force. Over half of the trauma is located on the posterior aspect of the crania, suggesting the victims were attacked from behind. Sixteen of the fractures observed were perimortem and two were antemortem. The distribution and characteristics of the fractures suggest that some of the Tripolye individuals buried at Verteba Cave were victims of a lethal surprise attack. Resources were limited due to population growth and migration, leading to conflict over resource access. It is hypothesized that during this time of change burial in this cave aided in development of identity and ownership of the local territory.
Ancient DNA studies have established that Neolithic European populations were descended from Anatolian migrants who received a limited amount of admixture from resident hunter-gatherers. Many open questions remain, however, about the spatial and temporal dynamics of population interactions and admixture during the Neolithic period. Here we investigate the population dynamics of Neolithization across Europe using a high-resolution genome-wide ancient DNA dataset with a total of 180 samples, of which 130 are newly reported here, from the Neolithic and Chalcolithic periods of Hungary (6000–2900 BC, n = 100), Germany (5500–3000 BC, n = 42) and Spain (5500–2200 BC, n = 38). We find that genetic diversity was shaped predominantly by local processes, with varied sources and proportions of hunter-gatherer ancestry among the three regions and through time. Admixture between groups with different ancestry profiles was pervasive and resulted in observable population transformation across almost all cultural transitions. Our results shed new light on the ways in which gene flow reshaped European populations throughout the Neolithic period and demonstrate the potential of time-series-based sampling and modelling approaches to elucidate multiple dimensions of historical population interactions.
There were some interesting finds on a regional level, with some late survival of hunter-gatherer ancestry (and Y-DNA haplogroups) in certain specific sites, but nothing especially surprising. This survival of HG ancestry and lineages in Iberia and other regions may be used to revive (yet again) the controversy over the origin of non-Indo-European languages of Europe attested in historical times, such as the only (non-Uralic) one surviving to this day, the Basque language.
This study kept confirming the absence of Y-DNA R1b-M269 subclades in Central Europe before the arrival of Yamna migrants, though, which offers strong reasons to reject the Indo-European from the west hypothesis.
Here are first the PCA of samples included in this paper, and then the PCA of ancient Eurasians (Mathieson et al. 2017) and modern populations (Lazaridis et al. 2014) for comparison of similar clusters:
Human ancestry can only help solve anthropological questions by using all anthropological disciplines involved. I have said that many times in this blog.
Correlation does not mean causation
Really, it does not.
You might think the tenet ‘correlation does not mean causation‘ must be evident at this point in Statistics, and it must also be for all those using statistical methods in their research. But it is sadly not so. A lot of researchers just look for correlation, and derive conclusions – without even an initial sound hypothesis to be contrasted… You can judge for yourself, e.g. reading the many instances of this complaint in recent publications of Biomedical and Social Sciences, on the interesting blog Statistical Modeling, Causal Inference, and Social Science.
In anthropological questions regarding Indo-European studies there is an added handicap: not taking correlation to mean causation does also mean – to avoid at least the most obvious confounders – taking into account the multiple linguistic and archaeological data that are available right now, to explain the expansion of Indo-European languages.
You might also believe that international researchers in Human Evolutionary Biology – after all, this is essentially a biomedical discipline – are acquainted with statistical methods and their problems when applied to their field. And that scientific journals – and especially those with the highest impact factors, like Nature, Science, or PNAS – have professional, careful reviewers who would never accept papers that equal correlation with causation, especially when Social Sciences are involved (because this alone might make errors grow exponentially…). Sadly, this is obviously not so, either.
Both studies [Haak et al. (2015) and this one] found a genetic affinity between samples from a central European culture known as Corded Ware, which existed from around 2500 bc, and samples from the earlier Yamnaya steppe culture. This similarity between distant populations is best explained by a substantial westward expansion of the Yamnaya or their close relatives into central Europe (Fig. 1b). Such an expansion is consistent with the steppe hypothesis, which argues that Corded Ware cultures were a conduit for the dispersal of Indo-European languages into Europe.
More interesting than these vague words – and the short, almost invisible suggestion that Yamna may not be exactly the population behind Corded Ware peoples – are the maps that illustrated in Nature their risky hypothesis: they called it “steppe hypothesis“, like that (in general terms), as if everyone defending a steppe origin for Proto-Indo-European would support such a model, when they actually referred to the specific hypothesis of one of their authors (Kristiansen), one of the few archaeologists who keep Gimbutas’ concept of the ‘Kurgan peoples’ alive, based on the Corded Ware culture:
In many publications that followed, the trend has been to reproduce this graphical model, by asserting (or implying) that Bell Beaker peoples were the result of subsequent Corded Ware migrations, and indeed that Corded Ware peoples migrated from the Yamna culture, and were thus the vector of expansion for Indo-European languages in Europe.
We shall see then just a rather surreptitious shift in terminology from ‘Yamnaya’ to ‘steppe’ component, to adapt to the new data – i.e. some damage control while the ship of ‘Yamnaya ancestry’ capsizes – but little else. “Earlier ‘Yamnaya ancestry’, you say? Just, you know, let’s call it ‘steppe ancestry’ and shift the expansion of Indo-European languages to one or two thousand years earlier, and done!”
The damage of this post-truth genetics is already done: we will see the unending distribution on the Internet in general, and on social networks in particular, of these grandiose conclusions, of far-fetched Indo-European migration models that include the Corded Ware culture, of simplistic maps with apparently harmless ‘arrows of migration’ (like the above) representing fictional population movements suggesting nonexistent dialectal branches.
You might be one of those sceptics wary of so many boring statistical rules: “But it’s a safe reasoning: Yamanaya samples have an ‘ancestral component’ that is found elevated in Corded Ware samples, and less so in Bell Beaker samples, and PCA showed a similar result…so the migration model Yamnaya -> Corded Ware -> Bell Beaker is a priori correct, right?”
The ‘Future American’ hypothesis
Let me illustrate this attractive “Correlation = Causation” argument, using it to solve the problem of Future American languages.
Suppose we live in a future post-apocalyptic world ca. 3500 AD, with no surviving historical records before 3000 AD. None. Just investigation of cultures and their relationship by Archaeology, proto-languages reconstructed and language families identified by Linguistics, etc.
We have thus Future Germanic and Future Romance as the only language families spoken in Future Western Europe and in the Future Americas, in a distribution similar to the present day*, and we have certain somehow related archaeologically-defined cultures on both sides of the Atlantic, like Briton, Iberian, Norman, or Lowlandish, although their distribution remains partly undefined in time and space.
* If you are really curious about this scenario, you can read about the potential evolution of a Future North-American language.
But what languages did the ancestors of Future Americans speak, and who spread them? That question remains far from being settled by our future researchers, in spite of the solidest linguistic and migration models (talking mainly about Briton and Iberian cultures): too many authorities out there questioning them, fighting to impose their own pet theories.
Suddenly, the newly developed field of Human Ancestry comes to save the day. So let’s say we have this map of ancient samples recovered (dated from, say, the 6th to the 18th century AD), and our study is centered on the newly described “Western European” component (a precise combination of, say, WHG+steppe), which peaks in early samples from the Low Lands – hence we call it, quite daringly, “Lowlandic component“.
Our group is keen to demonstrate that the ancient Lowlandic culture described in Archaeology (marked especially by the worldwide distribution of tulips among other traits) is the origin of Western European and American languages… Now, let’s reach conclusions about migrations in the Middle Ages!
PCA shows that South-West European samples cluster closely to some North-West European samples, and that some late South American samples available cluster at some distance from North American samples – nearer to a native component represented by two individuals with 0% Lowlandic ancestry and a different cluster in PCA. And some North-American samples cluster quite closely to North-West European samples.
Based on the decrease in ‘Lowlandic component’ in the different samples and on PCA, we conclude that Lowlandic peoples (“or their close relatives”) must have migrated at the same time to North America, South America (or potentially from North America to South America?) as well as western, central, and northern Europe. Both migration events must have happened roughly at the same time, in part because both distinct language families appear in a north-south distribution, and Proto-Lowlandic must be (according to Genetics) the ancestor of both, Proto-Future-Germanic and Proto-Future-Romance.
That makes a lot of sense! A huge Lowlandic pressure for migration, you see. Push-pull mechanisms and stuff. A Lowlandic Empire probably (scattered remains are found everywhere)! And, judging by the presence of the ‘Lowlandic component’ in Future East Europe from the Elbe to the Vistula, maybe Lowlandic peoples spread Proto-Slavic, too! We can even date the common Lowlandic-Slavic proto-language this way! So many groundbreaking conclusions!
Future scholars supporting the Lowlandic homeland are on fire; they can’t get enough of publishing papers on the subject. “Two different Future American language families with cultural origins in Britain and Iberia, my ass! Because genetics.”
And don’t forget the future people of haplogroup R1b-U106 and high Lowlandic component: Wow, they are the heirs of those who expanded Future Germanic and Future Romance languages everywhere, aren’t they? How proud they must be. And who wouldn’t want to have these tall, blond, blue-eyed Lowlanders as their forefathers? Personalised genetic analysis is selling like crazy: “let’s know our Lowlandic percentage!”. Everyone is happy, colourful maps with lots of arrows and shit…
But – your future you might ask in awe, seeing that this doesn’t sound quite right, based on your basic archaeological and linguistic knowledge:
What about specific models of migration proposed to date? The solidest ones, not just anyone that seems to fit?
What about the dialectal classification of languages? The mainstream ones, not those that are compatible with this interpretation?
What about archaeological cultures to which individual samples belonged?
What about the actual dates of each sample? And how this date relates to the state of the culture to which it belongs?
What about the haplogroups, and the actual subclade of each haplogroup?
What about the territories, cultures, and dates not sampled, could they change this interpretation in light of known archaeological models?
And what about the actual origin of that ancestral component they so frivolously named? Dit it really appear ex nihilo in the Low Lands, and expanded from it?
“Who cares! This new data is sooo coool… And it proves what we wanted, what a coincidence! And it’s numbers, mate! Numbers don’t lie.”