Refugees, Forerunners or Simply Uncommon? – J2b-L283 Paraclades in Sardinia and Tuscany

Two of the earliest branching points of the line of J2b-L283 from which most living men descend, J2b-Z600, have highest diversity in Sardinia and Tuscany. The ancient samples in Sardinia date to the Nuragic culture, predating the Phoenicians.

What Lazaridis et al. wrote in 2022

J-Z600, the parent node of J-Z585, includes four additional individuals, one of which is from Croatia, and three of which are from the Late Bronze Age Nuragic culture in the island of Sardinia, thus suggesting that this culture included individuals of Bronze Age Western Balkan origin (these might have Italian intermediaries, but we do not detect any J-Z600 in mainland Italy prior to the aforementioned Iron Age sample from Rome*).

The genetic history of the Southern Arc: A bridge between West Asia and Europe (Iosif Lazaridis, Songül Alpaslan-Roodenberg et al. 2022)

*There are now older J2b-Z600 found in mainland Italy prior to the Civitavecchia, Tuscany sample. You can see them on this map but I’m not going to mention them in this post. Instead I discuss some possible explanations for how these rare, early branching points of J2b-Z600 ended up in Sardinia / Etruria.

TMRCA and Geographic Distribution of the Paraclades

This part of the tree hasn’t changed in the last few years since the establishment of the European/Tunisian Jewish line or the sample from Lucca which formed a subclade with the scientific sample from Firenze.

J2b-Z2509 has two children, J2b-YP157 and J2b-Z585. The former is only found in Sardinia and has been there since at least the 12th or 13th century BCE based on the oldest sample ORC003. Neither YFull nor FTDNA provide a TMRCA estimate for J2b-YP157 because the samples are too low coverage (1-3x).

J2b-Z2509’s TMRCA is estimated at 3500-3100 BCE (these are estimates from FTDNA and YFull, not a confidence interval).

J2b-Z585’s TMRCA is estimated at 3100-3050 BCE (FTDNA and YFull estimates are very close). He sired four surviving lines. Three are very rare and one is the prolific J2b-Z615 that all other J2b-Z585 descend from.

The Rare Children of J2b-Z585 are:

  • J2b-BY100002, TMRCA 1300 CE, European/Tunisian Jewish line found at least in Tunisia, Sicily, Greece and Istanbul
  • J2b-FT289318, TMRCA 700 CE, only found in Tuscany
  • J2b-YP113, no TMRCA due to low resolution samples, only found in Sardinia

J2b-Z615 – Granddaddy of Everybody Else and MOK15

The ancestor of everyone else in this line lived about 3000-2950 BCE and sired a few subsequent lines at that time. His descendants started to proliferate again (i.e. siring more than a single surviving line) 7 SNPs later down the J2b-Z597 line, estimated at 500 years later.

Starting with J2b-Z597 diversification accelerates until about 4100-4000 BCE when a number of lineages start to sire subsequent lineages defined by a single SNP. The peak diversification rate during this time is still not as high as that achieved later during the Iron Age, but the lineages that formed later have a better chance of not having died out, so there is bias in doing a straight comparison.

Diversity computed by HRAS as branches formed per hundred year bin. There is a noticeable peak around 4100 BCE, though it is dwarfed by a later Iron Age peak.

We are lucky to have the ancient sample from Mokrin, Serbia, MOK15, that anchors J2b-Z615 to a location – the central Balkans. This one sample, that lived 1000 years after the MRCA of J2b-Z615, isn’t proof that the line originated in that exact place, but we know that some were living there by that time.

The Vučedol Culture

The Vučedol Culture flourished from 3000-2200 BCE and is named for the typesite Vučedol, located on the right bank of the Danube about midway along the Croatia-Serbia border. According to Wikipedia, the archaeological horizon “completely or partially covered 14 of today’s European countries – the Czech Republic, Slovakia, Austria, Hungary, Romania, Slovenia, Italy, Croatia, Bosnia and Herzegovina, Serbia, Montenegro, Kosovo, Albania and one settlement has even been registered in Eastern Greece”.

Per wikipedia, the main invention of this culture is considered to have been making tools from arsenical copper. This is why most of their earlier sites are not far from mountain ranges and the quest for copper deposits likely drove their expansion.

Also per wikipedia, “The settlement sites destroyed earlier eneolithic settlements, and new Vučedol settlements also developed in regions where none previously existed.”

The Y-DNA samples found to date include at least 4 samples, two from separate sites in Beli Manastir, NE Croatia confirmed R1b-Z2103, one from Çinamak, Albania who is R1b-M269 and a G2a sample from Croatia. So there is no ancient Y-DNA evidence that J2b-L283 was living within this culture.

However if J2b-L283 actually represented 20% of all men at the time, there would be a 0.8 ^ 4 = 41% chance that given four random ancient samples, we would not find a J2b-L283. This is very simple math that I recommend to use in other cases to add more objective and easy to understand facts on which theories can be based. It can help us get our heads around effects due to uneven sampling rates. In our case, two samples are from the exact same town so we may rather consider them as one data point. In that case, given three data points, there would be a 0.8 ^ 3 = 51% chance that no J2b-L283 is found if the true level was 20%. So as more samples are found, whatever their haplogroup is, we can still make incremental progress in assessing likelihood of a certain presence.

The recent paper Ancient DNA reveals the origins of the Albanians (June 2023) states that the earliest known sample with EHG (Eastern Hunter Gatherer) autosomal DNA in the western Balkans is I14689 from Çinamak, Albania. This sample is geographically very peripheral to Vučedol but there were Vučedol culture items in the grave. The authors posit that given his 70% steppe autosomal component and that he is found in the first kurgan burials in the Balkans, that his ancestors were likely recent arrivals from the steppe.

We’ll come back to this Balkan line of R1b-Z2103 a little later because our J2b-Z615 MOK15 was found in the same burial site in Mokrin as two men of R1b-Z2103>CTS7556. They belong to the Maros culture.

Ljubljana-Adriatic Style/Culture

Inconveniently, there is no Wikipedia article on the “Ljubljana Culture”.

However in the article on Vučedol Culture there is a regional variant described as “Slovenian – West Croatian (Ljubljana Marsh type)”.

Jake Marcisz kindly referred me to this article about pile dwellings and the Ljubljana culture. It is very dense reading with the pottery description language that I am unfamiliar with but I found something of interest for J2b-L283.

There are pile dwellings in the Ljubljana marshes similar to those found on both sides of the Alps, however their material culture resembles that found from the Karst Plateau to caves further south along the Adriatic coast. Leghissa considers the culture in the Karst to be part of the Ljubljansko barje tradition rather than the Adriatic tradition.

“It is Deschmann’s pile-dwelling sites in the Ljubljansko barje that likely represent the area of origin of the Ljubljana culture.”

However the final sentence states that the relationship between the Adriatic variant and the Ljubljana variant is not yet understood. The age-estimates actually indicate that the ‘Adriatic variant’ predates the Ljubljana proper culture, however the dating is unreliable.

“He uses stratigraphy, metal artefacts and radiocarbon dates to attribute the Ljubljana-Adriatic style to the first half of the 3rd millennium BC. Considering the dates for the Montenegrin tumuli, the Ljubljana-Adriatic style is contemporary with the Vučedol style, not only the Late, but also the Classic. In the mid-3rd millennium BC, the Cetina sytle is believed to have replaced the Ljubljana-Adriatic style.”

So the Cetina style replaced the Ljubljana-Adriatic style roughly during the time of accelerating diversification of subclades of J2b-Z615.

5000 Years Ago There May Have Been Five Clans of J2b-Z2509

At the time of the diversification of J2b-Z615 about 5000 years ago, there were only four other lines that we know of so far:

  • J2b-YP157
  • J2b-Z585>BY100002
  • J2b-Z585>FT289318
  • J2b-Z585>YP113

Five thousand years ago, each of the four lines may have been represented by a single chain (a man and his father and/or son), though it is likely that each may have been represented by some dozens of clan members whose lines later died out.

If we think about these different lines as clans, then including J2b-Z615 there were 5 “clans”.

The paraclades of J2b-Z2509 nearly died out and if they had originally been living on the Adriatic coast, they eventually moved away from there and/or died out there.

Theory: Five Clans of J2b-Z2509 Migrated to the Adriatic Coast of Croatia Following Rivers by 3000-2500 BCE

Whether there was (I) a migration of five clans to the Adriatic coast of Croatia by 3000-2500 BCE or whether (II) a single ancestor migrated there a few hundred years earlier and sired the lineages there, it doesn’t change the rest of the theory much going forward from that point.

The only difference it makes is regarding the direction from which the ‘inland’ J2b-Z615 clan’s immediate ancestors came from – either from further inland (I) or back from the coast (II).

The reason I pick the Adriatic coast as the place of origin or end of a migration of most surviving J2b-Z2509 is that I propose that they were “pushed out” of their Balkan homeland by their more numerous / successful cousins from inland, the bearers of the Cetina culture. I propose the area of the coast near where the Cetina culture originated because we already have a precedent of one culture coming to the Adriatic by that route.

There may have been a period of population growth in the core Cetina zone, the upper stream of the Cetina and lower Neretva, prior to the development and spread of their culture throughout the Adriatic, replacing the Ljubljana-Adriatic culture and then going further. The population growth in the core Cetina zone could have been due to the dissemination of new ideas and/or technology ultimately transmitted from groups traveling up or down the Danube.

Galaty et al. (2015) argue that a ‘warrior culture’ including “ideas related to warrior aristocracy” spread from Europe to Greece through contact with the Cetina culture, along with the tradition of tumulus burial.

So the initial growth of J2b-Z615 may have been simply having more children with their own women due to favorable conditions, while the accelerating growth could in part be attributed to the replacement of other male lines as a result of successful military endeavors / raids.

For instance, J2b-Z615, with their militarily superior culture, may have gradually taken control of the areas along the Adriatic inhabited by the four “clans” of their cousins.

In that case, there may have been real pressure to leave the hostile environment.

If the J2b-Z2509 paraclades had been living on the Adriatic coast for some time before being displaced by J2b-Z615, they may have had enough time to develop a maritime lifestyle before their J2b-Z615 cousins did. The quickest way to develop the needed skills would be through learning from the indigenous peoples.

Flight of Ljubljana Adriatic J2b-Z2509 Paraclades to the Ljubljana Marsh?

The article about the pottery of the Ljubljana cultures (Ljubljana proper and Adriatic ‘variant’) raises an interesting possibility regarding how the J2b-Z2509 paraclades may have ultimately migrated to Sardinia or at least Etruria.

Leghissa states that Franco Nicolis and other researchers think that the influences for the “barbed wire” Bell Beaker, very common in northern Italy and southeastern France from 2150 – 1900 BCE, probably came from the Ljubljana culture.

If the unreliable dating methods are correct, and the Ljubljana proper culture was introduced to that area after it had been originally developed in the Ljubljana-Adriatic zone, then perhaps it was the J2b-Z2509 paraclades who brought the culture with them to the Ljubljana marshes as they sought refuge from their more warlike Cetina culture cousins.

Later the J2b-Z2509 may have migrated further west to the Karst, into northern Italy or even southern France, bringing the “barbed wire” Bell Beaker style with them, though this line hasn’t yet been found in France.

J2b-Z2509 Paraclades as “Forerunners” of Cetina in the Adriatic / Ionian / Sicily Seas?

If, instead, J2b-Z2509 had sailed south along the Adriatic to the Ionian and Sicilian Seas, their ultimate migration to Sardinia and Etruria could have come from south via the Tyrhennian Sea.

In that case, the paraclades may have represented the earliest migrations along the same sea routes that the Cetina culture would soon traverse and dominate. The material culture of the “forerunners” may have been Ljubljana-Adriatic, fully-developed Cetina or some transitional phase.

Since the discovery of ancient 6th century BCE J2b-L283>Z38240 samples in Kerkouane, an ancient Phoenician settlement in northeast Tunisia, we know that lineages attested by Bronze Age samples from the western Balkans were able to migrate to the Tyrhennian Sea by way of the Adriatic and Ionian Seas by the Iron Age.

The ancient J2b-Z2509 on Sardinia predates the ancient samples from Kerkouane by 600 or 700 years and is a different line of J2b-L283. However I don’t think it is a big stretch to assume that the migration of J2b-Z2509 to Sardinia could have come from the south. After all, the single site where they are found is in Perdasdefogu, the southeastern part of the island.

To add: Autosomal info about ancient Sardinian samples

While I do not find any literature mentioning Cetina culture finds on Sardinia, genes do not always equal pottery just like they do not always equal language or ethnic identity. Another possible reason for lack of Cetina finds is that the J2b-Z2509 migration may have preceded the development of that culture, as I posited earlier.

Or Were J2b-Z2509 Paraclades Simply Rare Lines from the Adriatic Coast of Croatia that Later Accompanied Cetina Cultural contacts to Sicily and northern Africa?

If a line exists in Sardinia and in other places, then if this line is very rare, we are much more likely to first find it in Sardinia, sampled at more than 20 times the per capita rate of the rest of Italy that has not been targeted by scientific studies*.

While Liguria and Friuli-Venezia-Giulia are sampled slightly more than the average for Italy, the remaining core regions of Northern Italy such as Veneto, Lombardia, Emilia-Romagna, Piemonte are sampled 1.8-3.3 times less than the rest of Italy*.

*excluding Sardinia and Tuscany because these regions were the focus of targeted academic testing

However, because there are other rare lines in J2b-Z597 that are NOT found in Sardinia, I think the oversampling of Sardinia is not on its own enough to explain the relatively high diversity of J2b-Z2509 there if they were simply a rare line within the Cetina proliferation. So I prefer the aspects of the previous theories that emphasize an origin on the Croatian coast of the Adriatic vs more inland and/or a relatively early eviction from the Balkans that provide some additional explanation.

J2b-Z615 and R1b-Z2103>CTS7556 Buried in the same Cemetery in 1900 BCE in Mokrin, Serbia

Add: link to Maros paper

The fact that these samples were buried together could reinforce the idea that this specific branch of J2b-Z615 was the one with closer contacts to Yamnya-related groups originating from the steppe.

Given that in the Southern Arc paper, they posited that Indo-European languages (or “Indo-Hittite” as it were) may have originally derived from groups in the Caucasus (because genetically this is the main thing that links Hittites with the peoples from the steppe), I think it is important now to decouple the simplification that speakers of Indo-European languages = migrants from Steppe.

Given that the oldest ancient J2b samples come from the South Caucasus and West Zagros mountains, where Laziridis posits are the potential roots of “Indo-Hittite”, the ancient J2b-L283 may have already been speaking an Indo-European or “Indo-Hittite” language of some sort prior to any contact with R1b-Z2103 who would have been speaking a “fully-formed” Indo-European language from the steppe.

Explainer: if Indo-Hittite is the common ancestor of the language families, then I think “Indo-European” should refer to the languages that developed later in the steppe. For more information, I found this Quora answer useful –

So the contact between J2b-Z615 and Yamnaya-related R1b-Z2103 in Mokrin and elsewhere in Europe may have introduced steppe-origin military and other cultural traditions to J2b-Z615 but they may have been speaking somewhat related languages already – in fact, language similarity may have facilitated their coexistence.


Unfortunately there are some uncertainties that make it hard to resolve the theories at the moment. The Sardinian ancient and modern samples have lower coverage (1-3x) that makes it impossible to reliably estimate their MRCAs.

Another issue is that most of Northern Italy is sampled 1.8 to 3.3 times less than the rest of Italy. Also very few ancient DNA from the Bronze Age.

The other uncertainty is regarding the relationship between the Ljubljana proper and Ljubljana-Adriatic, i.e. which culture originated from the other, if that happened?

I’m also not sure what conclusions can be drawn from the sample from Moldova that was not published together with the Southern Arc.

These posts are the opinion of Hunter Provyn, a haplogroup researcher in J-M241 and J-M102.

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