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Uniparental analysis of Deep Maniot Greeks reveals genetic continuity from the pre-Medieval era

It’s not that they aren’t autosomally Greek; rather, they represent the segment of J2>L26 that migrated from the Caucasus region (Maikop, Kura-Araxes, or nearby) to the Aegean between 4000–2000 BC and did not mix with other populations high in EEF, unlike the rest of the Greeks. By retaining over 50–80% of their original J2-L26> haplogroup, instead of shifting in the PCA, they effectively drew other populations toward their cline, placing them close to modern Caucasians.

I’m not an expert on J2 but this haplogroup in Deep Maniots does not need a Caucasus to Greece back-and-forth explanation.
Ancient DNA already shows that J2 was present in Anatolia, the Aegean, and the Near East since the Neolithic. Aegean populations were not peripheral to this world, they were central to it. So J2 in Greece was part of the same long-standing eastern Mediterranean continuum.
Deep Maniots are overwhelmingly J2, especially a narrow subset of J2a. Chechens and Ingush are also heavily J2, again dominated by a few deep-rooted Caucasus-specific branches.
In the NMDS plot populations dominated by the same haplogroup cluster together, even if they split 6,000-10,000 years ago.
Deep Maniots land between Chechens and Ingush because they share deep J2 Y-DNA ancestry, not because they are part of the Caucasus-specific expansions.
The plot does not imply that Deep Maniots came from the Caucasus, it implies that J2 has multiple ancient regional branches, and isolated populations dominated by J2 can cluster together in a STR space even if their divergence is very old.​
 
I’m not an expert on J2 but this haplogroup in Deep Maniots does not need a Caucasus to Greece back-and-forth explanation.
Ancient DNA already shows that J2 was present in Anatolia, the Aegean, and the Near East since the Neolithic. Aegean populations were not peripheral to this world, they were central to it. So J2 in Greece was part of the same long-standing eastern Mediterranean continuum.
Deep Maniots are overwhelmingly J2, especially a narrow subset of J2a. Chechens and Ingush are also heavily J2, again dominated by a few deep-rooted Caucasus-specific branches.
In the NMDS plot populations dominated by the same haplogroup cluster together, even if they split 6,000-10,000 years ago.
Deep Maniots land between Chechens and Ingush because they share deep J2 Y-DNA ancestry, not because they are part of the Caucasus-specific expansions.
The plot does not imply that Deep Maniots came from the Caucasus, it implies that J2 has multiple ancient regional branches, and isolated populations dominated by J2 can cluster together in a STR space even if their divergence is very old.​

MMDS projections do not drastically change the overall positioning compared to an autosomal PCA. The Portuguese, Spanish, Basques, and Italians are distributed very similarly to an autosomal PCA because both populations share more than 50% of ZZ11>, but Sephardic Jews, who autosomally position near Eastern Mediterraneans in this STR PCA, are located between Italians and Spaniards. This occurs because R1b lineages have more compact STRs than any J1, J2, or E haplogroups.

In other words, 30% of J1 with all branches converging in P58> around 8000 BC is not the same as 10% converging in all ZZ11> around 3000 BC. ZZ11 will carry more weight than JP58.

They measure different things, and it’s important to understand what real relationship they reflect.

For practical purposes, it is almost the same as looking at an autosomal PCA but removing the autosomal SNPs contributed by the mitochondrial component, which can shift some populations’ positions slightly, yet in practice most populations are distributed very similarly.

The problem is that you “create” or “distort” mixtures that never actually existed.

This type of mapping and filtering helps with terminal Y-SNP refinements.

Hypothetical example:

At a time when ZZ11> SNPs were not yet cataloged, let’s say in the early 2000s, they start unearthing Bell Beakers.

Bronze Age ZZ11> individuals would be autosomally indistinguishable in an autosomal PCA, but an STR PCA would reveal two clines that would later correspond to DF27 and U152.

A DF27 with 7 out of 8 great-great-grandparents U152 would autosomally appear closer to U152.

A U152 with 7 out of 8 great-great-grandparents DF27 would autosomally appear closer to DF27.

But after applying STR filtering, each would appear in its original place.

Rare cases, like the Maniot Greeks, should ideally be projected using:
  1. Autosomal PCA
  2. Y-STR PCA
  3. Mt-STR PCA
All three together to clearly determine the origin of each component and understand the sequence in which IBD segments were shared.

The problem with STR projections is that they mix male-exclusive data, which is impossible in reality. That’s why “strange” things appear, such as 10% of ZZ11 pulling Jews toward the Western-dominated zone. They also depend heavily on the populations selected.

Therefore, there are two types of populations:
  • Those with 5000 years of autosomal (cultural) and 5000 years of ethnic (STR) history, like Portuguese, Basques, Spaniards, and Italians.
  • Those with 4000 years of autosomal (cultural) history, like Jews, Maniots, or Romani, but whose ethnic Y-STR makeup is a much more recent mixture or creation.
A culture with interrupted ethnic continuity is not the same as a marginal culture gradually absorbing other ethnicities.

For example, the English have 5000 years of autosomal history, but only 1500 years of STR continuity from the P312-U106 mixture.

STR mappings are a quick way to see differentiations without refining terminal SNPs one by one. If you already know the deep SNPs from the start, you don’t need STR filtering because you already know how lines differ.

The cost difference: STR filtering for 100 samples would be around €1000, while terminal SNP refinement would be €1000 per sample.

In the specific case of the Maniot Greeks:
  • 4000 years culturally and autosomally Greek
  • 2700 years Y-STR continuity from Mycenaean + Classical Greeks
  • 1300 years Y-STR continuity of isolated Greeks, which enhances their Caucasian ancestry







In this case, there are no populations coming from elsewhere, only internal genetic drift.
 
MMDS projections do not drastically change the overall positioning compared to an autosomal PCA. The Portuguese, Spanish, Basques, and Italians are distributed very similarly to an autosomal PCA because both populations share more than 50% of ZZ11>, but Sephardic Jews, who autosomally position near Eastern Mediterraneans in this STR PCA, are located between Italians and Spaniards. This occurs because R1b lineages have more compact STRs than any J1, J2, or E haplogroups.

In other words, 30% of J1 with all branches converging in P58> around 8000 BC is not the same as 10% converging in all ZZ11> around 3000 BC. ZZ11 will carry more weight than JP58.

They measure different things, and it’s important to understand what real relationship they reflect.

For practical purposes, it is almost the same as looking at an autosomal PCA but removing the autosomal SNPs contributed by the mitochondrial component, which can shift some populations’ positions slightly, yet in practice most populations are distributed very similarly.

The problem is that you “create” or “distort” mixtures that never actually existed.

This type of mapping and filtering helps with terminal Y-SNP refinements.

Hypothetical example:

At a time when ZZ11> SNPs were not yet cataloged, let’s say in the early 2000s, they start unearthing Bell Beakers.

Bronze Age ZZ11> individuals would be autosomally indistinguishable in an autosomal PCA, but an STR PCA would reveal two clines that would later correspond to DF27 and U152.

A DF27 with 7 out of 8 great-great-grandparents U152 would autosomally appear closer to U152.

A U152 with 7 out of 8 great-great-grandparents DF27 would autosomally appear closer to DF27.

But after applying STR filtering, each would appear in its original place.

Rare cases, like the Maniot Greeks, should ideally be projected using:
  1. Autosomal PCA
  2. Y-STR PCA
  3. Mt-STR PCA
All three together to clearly determine the origin of each component and understand the sequence in which IBD segments were shared.

The problem with STR projections is that they mix male-exclusive data, which is impossible in reality. That’s why “strange” things appear, such as 10% of ZZ11 pulling Jews toward the Western-dominated zone. They also depend heavily on the populations selected.

Therefore, there are two types of populations:
  • Those with 5000 years of autosomal (cultural) and 5000 years of ethnic (STR) history, like Portuguese, Basques, Spaniards, and Italians.
  • Those with 4000 years of autosomal (cultural) history, like Jews, Maniots, or Romani, but whose ethnic Y-STR makeup is a much more recent mixture or creation.
A culture with interrupted ethnic continuity is not the same as a marginal culture gradually absorbing other ethnicities.

For example, the English have 5000 years of autosomal history, but only 1500 years of STR continuity from the P312-U106 mixture.

STR mappings are a quick way to see differentiations without refining terminal SNPs one by one. If you already know the deep SNPs from the start, you don’t need STR filtering because you already know how lines differ.

The cost difference: STR filtering for 100 samples would be around €1000, while terminal SNP refinement would be €1000 per sample.

In the specific case of the Maniot Greeks:
  • 4000 years culturally and autosomally Greek
  • 2700 years Y-STR continuity from Mycenaean + Classical Greeks
  • 1300 years Y-STR continuity of isolated Greeks, which enhances their Caucasian ancestry
In this case, there are no populations coming from elsewhere, only internal genetic drift.

If I understand correctly, you are saying that Y-STR NMDS plots is a weakened or filtered version of autosomal PCA. But the two methods measure entirely different things. Autosomal PCA reflects genome-wide ancestry shaped by recombination and admixture across both sexes. Y-STR NMDS reflects only a single non-recombining paternal lineage and is dominated by drift, founder effects, mutation-rate variance, and sampling structure. Any visual similarity between the two is incidental and occurs only where male-line history happens to follow the same broad geographic patterns as overall ancestry. This similarity breaks down immediately in cases of isolation, endogamy, or sex-biased history, which is why groups like Maniots or Jews look normal in autosomal analyses but “odd” in STR plots.

The idea that certain haplogroups “carry more weight” or that small percentages of lineages can “pull” populations misunderstand how distance-based ordinations work. STR NMDS does not weight lineages by age, frequency, or autosomal contribution. Apparent influence comes from homogeneity, drift, or overrepresentation in the sample, not from real ancestry proportions. These are mathematical artifacts, not biological signals.

STR plots cannot undo autosomal mixing or recover a population’s “original” position. They remove information rather than refine it by ignoring nearly all ancestry outside the paternal line.

Concepts like “ethnic Y-STR makeup” or “autosomal (cultural) history” are not meaningful in population genetics. Y-STRs describe paternal lineage structure, not ethnicity.

The idea that populations like Portuguese, Basques, Spaniards, or Italians have “5000 years of ethnic (STR) continuity” is not correct. Their Y-chromosome histories include multiple turnovers, bottlenecks, and expansions, especially during the Bronze Age and later periods. The Basques, in particular, are a counterexample, they have strong autosomal continuity relative to Neolithic farmers, but near-complete Y-chromosome replacement by Bronze Age R1b lineages. That is the opposite of uninterrupted “ethnic STR continuity.”

The arguments about Mycenaean continuity, enhanced Caucasian ancestry, or ethnogenesis cannot be inferred from Y-STR alone. Such conclusions require ancient DNA, dated Y-SNP phylogenies, and demographic modeling. At most, the Y-STR NMDS plot shows that Deep Maniots are a highly drifted, J2-dominated paternal population clustering with other J2-heavy, drifted groups in STR space, nothing more.​
 
Probably because they are Anatolians? They plot like Anatolians; they have the same genetic makeup as Anatolians. What else could they be? Greeks Islanders? They certainly aren't mainland Greeks, which would suggest a complete population replacement between 30 BC and 476 CE as the bulk of the Hellenistic samples plot with their predecessors.

They are Greco-Anatolians. People who had long since been Hellenized and considered proper Greeks but carried a slightly more Eastern/Caucasian heavy profile relative to the Greeks of the bronze age. I doubt it was a fast replacement but more realistically hundreds of years of gradual genetic overturn through migration, assimilation and fertility bias.

Or they need heavy Caucasian admixture to plot where they do, but Caucasian profiles in the Greek mainland are so far non-existent.

I agree that pure Caucasian proper profiles would have been quite rare, but your own image shows two Caucasian profiles, to be fair. Impossible to tell what time frame they are dated to of course, so we can't say whether ancient or medieval.

Imagine archaeologists from 2000 years in the future discovering the family graves of Pontic Greeks in Thessaloniki from the 20th century and proclaiming they were representative of the genetic profile of mainland Greeks. Lastly, the late Hellenistic sample is also supportive of the findings of the unleased paper that Greece began experiencing increased Steppe inflow from the Hellenistic to Roman transition.

I don't think Tenea profiles will be particularly uncommon for Peloponneisan and more broadly southern Greece by the Roman era. I think instead more northerly cities like Thessaloniki and also more broadly Macedonia will plot more like modern Greek Mainlanders due to retaining a much older, likely bronze aged more steppe heavy profile that is similar to what we see in Paeonia. There will almost certainly be an ancient north-south cline, just like what we are slowly finding in Italy. Modern Greeks have seen a reduction in this Anatolian like profile in favor of Northern Greek ancestry by comparison, implying a north to south fertility and migration bias throughout the middle ages.

Thanks for posting this picture however. It implicates strongly much of what I've long suspected.
 
I was more talking about the chronology of the split between the J-L930, I know about the influx, although I do not see much evidence to call it constant before the Hellenistic era. I had misread the part where the paper discussed the related haplogroups to it and misunderstood it as chronologically as spliting with the caucasus branches in the late bronze age when all the paper says it that it has related branches in the bronze ages but probably originates in the neolithic. If it had migrated out of anatolia in the late bronze age it would put a variable on the slight increase of steppe ancestry during the Greek dark ages found in Skourtanioti 2023 and would imply a bigger influx of steppe admixed individuals than initially thought by the increase.

Skourtanioti found continual increases of both steppe and Anatolian ancestry throughout the full span of the bronze age at the expense of local EEF ancestry which itself was of course derived nearly entirely from Anatolia during the neolithic. These are the factors I refer to when I say perpetual overturn. Ancestry from Anatolia gradually migrated and replaced what was previously dominant in the west. This pattern necessarily ends in Greece sometime prior to the conquest of the Turks.
 
This plot is not a Principal Component Analysis (PCA), it is a Non-Metric Multidimensional Scaling (NMDS) used for measure genetic distances from Y-chromosome STR markers. It is designed as a descriptive, exploratory visualization tool for patrilineal similarity and structure, and not as a measure of population ancestry, ethnicity, or autosomal relatedness. Is useful for studying male demographic history, and completely inappropriate for inferring autosomal ancestry or population affinity.
This plot is a distance metric originally designed for allele frequency differences, when applied to Y-STRs, it quantifies how different male lineages are, it does not model admixture. Is useful for exploring patrilineal structure, detecting clusters of related Y lineages, identifying founder effects or clan expansions and seeing whether certain male lines are shared across regions.
We can answer several questions like: are two populations sharing many Y-STR lineages, are they dominated by one or two paternal founders, is there high or low patrilineal diversity?
It’s a map of male-lineage similarity, distorted by mutation rate, drift, and sampling, useful for spotting patterns, but meaningless for measuring ancestry. We cannot say in a plot like this that if “Population A clusters with Population B they’re closely related genetically”, this is incorrect for this type of plot. The correct statement would be, “Some paternal lineages found in Population A resemble those in Population B.” Researchers still use it because when used properly, it can reveal hidden founder events and explain Y-haplogroup frequencies. It’s not a replacement to autosomal PCA / ADMIXTURE.
KEEKxG6.png

Apologies for the misnomer of this being a PCA. I agree with what you say; the NMDS may be useful for seeing how closely related an extremely small part of one's genetic expression is relative to norms of other populations, but as we know one must instead look at ancestral markers found on all chromosomes to quantify relatedness between ethnic or racial groups.
 
I agree with other posters that Greeks are one of the least tested populations both autosomally and haplogroups-wise. The Greek diaspora is much better tested than the Greeks in Greece. They tend to concentrate in the US, Germany and Australia with different geographic areas in Greece contributing to the genetic mix in each of those countries. For example the majority of Australian and American Greeks emigrated from the Peloponnesus and Central Greece with the majority of German Greeks emigrated from Northern Greece. I don't have any kind of insight on the composition of Greeks in the UK except that there are a lot of Greek Cypriots. So we have to be careful about any conclusions about the Greeks in general when it is the Greek diaspora that is getting tested and not Greece proper. There is no uniformity in the testing. For example, I have a lot of "cousins" from my mother's village getting tested whereas only one cousin from my father's village got tested. The two villages are 2 miles apart. People from my mother's village emigrated to Germany in much greater numbers than my father's village because they were poorer. I only know that they took the autosomal test and have no idea if any underwent Y-DNA testing.
 
If I understand correctly, you are saying that Y-STR NMDS plots is a weakened or filtered version of autosomal PCA. But the two methods measure entirely different things. Autosomal PCA reflects genome-wide ancestry shaped by recombination and admixture across both sexes. Y-STR NMDS reflects only a single non-recombining paternal lineage and is dominated by drift, founder effects, mutation-rate variance, and sampling structure. Any visual similarity between the two is incidental and occurs only where male-line history happens to follow the same broad geographic patterns as overall ancestry. This similarity breaks down immediately in cases of isolation, endogamy, or sex-biased history, which is why groups like Maniots or Jews look normal in autosomal analyses but “odd” in STR plots.

The idea that certain haplogroups “carry more weight” or that small percentages of lineages can “pull” populations misunderstand how distance-based ordinations work. STR NMDS does not weight lineages by age, frequency, or autosomal contribution. Apparent influence comes from homogeneity, drift, or overrepresentation in the sample, not from real ancestry proportions. These are mathematical artifacts, not biological signals.

STR plots cannot undo autosomal mixing or recover a population’s “original” position. They remove information rather than refine it by ignoring nearly all ancestry outside the paternal line.

Concepts like “ethnic Y-STR makeup” or “autosomal (cultural) history” are not meaningful in population genetics. Y-STRs describe paternal lineage structure, not ethnicity.

The idea that populations like Portuguese, Basques, Spaniards, or Italians have “5000 years of ethnic (STR) continuity” is not correct. Their Y-chromosome histories include multiple turnovers, bottlenecks, and expansions, especially during the Bronze Age and later periods. The Basques, in particular, are a counterexample, they have strong autosomal continuity relative to Neolithic farmers, but near-complete Y-chromosome replacement by Bronze Age R1b lineages. That is the opposite of uninterrupted “ethnic STR continuity.”

The arguments about Mycenaean continuity, enhanced Caucasian ancestry, or ethnogenesis cannot be inferred from Y-STR alone. Such conclusions require ancient DNA, dated Y-SNP phylogenies, and demographic modeling. At most, the Y-STR NMDS plot shows that Deep Maniots are a highly drifted, J2-dominated paternal population clustering with other J2-heavy, drifted groups in STR space, nothing more.​
So, what they did by mixing all the Y-STRs makes absolutely no sense when there is already a terminal SNP like J2>L26>L923.

STR mixes should only be done within the same SNP clade, not mixing J1 with J2 and R1b as they did in this case.

By simply looking at the SNP, the date, and the bottleneck size, you can already deduce what happened.

Since they mixed clades, the projection ends up resembling, to some degree, an autosomal PCA because different SNP clades were combined. That’s why I say it looks similar to an autosomal PCA, but without the autosomal SNP IBD segments that mtDNA contributes; they have nothing to do with each other since they’re only measuring, so to speak, the last ~1,000 years of Y-SNP variation.

NMDS STR graphs are only useful as an illustrative narrative to visually show recent patterns or anomalies, but if you understand what an STR and an SNP are, these kinds of projections are unnecessary.

STR PCAs are “confirmatory.”

Autosomal PCAs are “representative” based on pre-selected SNP sets; that’s why a sample with 5% coverage gives the same result as a WGS.

Y-17 STRs only reliably reflect roughly the last ~1,000 years.

Each SNP is a solid time marker; STRs show variation within that SNP but cannot be traced back thousands of years without direct archaeological evidence.

This is why mixing STRs from different lineages without considering the terminal SNP produces “absurd” results.

Example of misinterpretation:

In NMDS or PCA STR analyses, you see Sephardic Jews clustering between Spaniards and Italians, even though genetically they barely share ZZ11.

This is because:

• ZZ11> is very common in Spain and Italy (>50%), whereas Jews have very little (<10%).

• Each ZZ11 reaching Jewish populations comes from different lineages depending on the region, while P58 (representative of Jews) is much smaller and therefore has less STR variability.

Thus, STRs reflect recent homogeneity, not real ancestry or deep continuity.

This is why mixing all STRs from different SNPs pulls them toward the Western cline.


About the Basques and what I meant when I said their STR profile hasn’t “changed” but “evolved” over the last 5,000 years:

Considering SNPs and approximate dates:

• P312> 3500–3000 BCE → 90%

• P312>DF27> 2800–2700 BCE → 70%

• P312>L21> 2700–2600 BCE → 20%

• P312>DF27>Z195 2650 BCE → 40%

• P312>DF27>ZZ12 2650 BCE → 30%

• G2> pre-5000 BCE → 5%

• I2> pre-8000 BCE → 5%

STRs change slowly (~1,000 years per significant change). That’s why, if you take an average of Y-17 STRs over the last 1,000 years, they appear relatively homogeneous in a population with such SNP continuity.

This structure confirms 5,000 years of continuity because what certifies continuity are the SNPs, validated with ancient samples: EHU002, of non-steppe and non-Central European origin, confirmed by strontium analysis (P312>DF27>ZZ12 from 2600 BCE), shares direct continuity with modern ZZ12 in the Iberian Peninsula. While its exact STR profile isn’t identical, using multiple ancient specimens spaced every 500 years can demonstrate SNP continuity and also certify STR continuity.

P312> did not arrive from any external source; it is an internal Atlantic/Mediterranean expansion.

The correct method is to use SNPs as a base and STRs to analyze millennial evolution between consecutive SNPs. This allows reconstruction of 10,000 years of genetic history step by step, always validated with ancient samples.

In the autosomal context:

If you take any Iberian individual (Lusitania, Tartessos, Cogotas, Gallaecia, Asturias, or Argar), whether I2 from the Chalcolithic or P312 from Chalcolithic/Bronze Age, you will see:

• 60% Atlantic/Mediterranean ancestry, and

• From 2000–1000 BCE onward, ~40% derived from minor absorption of Amazigh, Phoenicians, Greeks, Romans, and Goths.

The Basques only absorbed ~10% foreign input.

Internal drift is minimal: over 5,000 years it does not drastically alter the autosomal landscape.

NMDS analyses of Y-STRs are only illustrative of recent male structure.

They cannot replace or refine autosomal DNA or deep lineage history.

In both types of PCA, Basques would appear in the same relative position to other groups because they have had very few changes and very high continuity. Portuguese, Italians, and Spaniards would also meet similar criteria.

Any attempt to extrapolate “ethnic continuity” or “cultural ancestry” from STRs alone is mathematically and biologically incorrect; SNPs and ancient samples of related lineages are required.

To interpret where this STR mixture projects, you must consider the SNPs used to generate that “projection mix.”

This explains the “pull” of Sephardic Jews toward the Western cluster or the “pull” of Maniots toward the Chechen STR cline.

Their positional fit in the Caucasian region in PCA is not coincidental; it is causal, because Y-STRs of J2 originated in the Caucasus, so they will always tend to reflect that cline, since that’s where the most basal J2 lineages exist.

Thus, seemingly unrelated phenomena are in fact related.

Both Maniots and Sephardic Jews appear in “odd” positions compared to an autosomal PCA, but for two completely different reasons: because different SNPs of different haplogroups are being mixed in the same NMDS STR.

In this case, the STR projection resembles autosomal PCA because of STR mixing:

For Jews, it is due to haplogroup mixing.

For Maniots, it is due to isolation.
 
This is empirically false.
They change drastically in projections that use only SNPs from a single Y lineage, such as J2-L26 and only derived clades.

In an STR-based PCA where different haplogroups such as E, J, I, or R are mixed, the result will resemble an autosomal PCA.

This is the case in this study.

Option 1 is used in biology to separate animal clades.

Option 2 is a representative mixture of Y lineages
 
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