A first, and intriguing glimpse at TRB West Group aDNA

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A good month ago, I presented here a new publication by Krause-Kyora e.a. “Neolithic and medieval virus genomes reveal complex evolution of Hepatitis B” that includes some interesting fresh aDNA from Central Europe. The most fascinating sample they provide is TRB-Tiefstichkeramik, Hildesheim-Sorsum, 3335–3107 cal BC, which closes an important geographical, chronological and cultural gap, and holds several surprises. Before presenting and discussing that sample, however, let me start with some background information:

TRB Groups acc. to Bakker (1989)

What means “TRB-Tiefstichkeramik”?

Most of you will know the “classic” structuring of TRB into North, West, East, South and Southeast Groups as first proposed by J.A. Bakker in 1989.  “Tiefstichkeramik” isn’t part of that structure, isn’t it? Well, in fact, it is: The little “A” in Bakker’s map, in-between the N, E, S and W Groups stands for the “Altmark Group” that in German is known as “Altmärkische Tiefstichkeramik“. However, Hildesheim-Sorsum isn’t in the Altmark, but a good 100 km further SW, in an area that would commonly be regarded as southern outskirt of the TRB West Group.

Then, I realised that “Tiefstichkeramik” was traditionally used as alternative label of the TRB West Group. This label is now widely regarded as obsolete, as (i) it only characterises a specific time horizon (ca. 3350-3000 BC) of TRB West, and (ii) “Tiefstich” decoration (“stab-and-drag” in English, partly with white fill) wasn’t  just characteristic for the Altmark and TRB West, but between 3500 – 3000 BC also dominated in TRB North, occasionally occured in the S and SE Groups, and, in its final blossoming with the Bernburg Culture, made it as far south as Franconia and Bohemia.

Tiefstich Beaker from Drenthe, NL (Wikimedia)
Amphora, Stettin Museum 1936 (Wikimedia)
Tiefstich sherd, Karleby, Västergötland
(Wikimedia)

Anybody interested in further details is referred to J.A. Bakker “The TRB West Group: Studies in the Chronology and Geography of the Makers of Hunebeds and Tiefstich Pottery”. That title  reveals the main feature of Tiefstich pottery: It is closely linked to the spread of megalithic burials (hunebeds in Dutch). In fact, TRB household pottery used to be rather plain and undecorated. The richly incised beakers were reserved for ritual purposes, and almost all of them were found in megalithic burials. So, unless a Dolmen or chamber grave is fully dismantled to collect organic residue between and below the groundstones for AMS dating, the best method for age determination is still analysis of the (Tiefstich) pottery it contains against fine-tuned regional chronologies (e.g. Brindley I-VII in the Netherlands).

The genesis of Nordic Megalithism is yet somewhat obscure. While the epicentre clearly lies on the Danish Isles, the earliest AMS dates come from the periphery: Borgstedt (n. Rendsburg) ca. 3960 – 3740 calBC, Albersdorf and Lüdelsen 36th cBC, Gökhem 17  3638–3378 calBC, Gotland 3619- –3351 calBC. The earliest safely dated Dolmen on Zealand (Ølstykke) is from the 35th cBC, contemporary with early megalithic burials on the Mecklenburg coast and on Rügen. Obviously, open waters didn’t inhibit the concept’s spread, so it may well have arrived from Brittany via Britain.

Available AMS dates for TRB enclosures show the same pattern: The recently discovered Hamremoen enclosure in SE Norway has yielded a surprisingly early dating to the 39th cBC; in the 37th cBC follows Büdelsdorf, just next to the a/m Borgstedt Dolmen.  Danish enclosures date to the late 36th cBC at earliest, and four of the eight early Danish enclosures cluster in N. Jutland.

Whatever the origin, from the 35th cBC onwards Megalithicism gained an enormous dynamic. By the late 34th century, it had reached East Frisia, and Drenthe in the NE Netherlands (the concentration of Helgoland flint in Drenthe hints at a possibly arrival by boat), as well as Scania and Central Mecklenburg. The spreading pattern through Lower Saxony has remained somewhat unclear, at least to me. In any case, the AMS dating supplied with the Sorsum sample indicates that the location was reached by the early 32nd cBC at latest.

Map of Red Helgoland Flint artefact finds

The expansion is best documented for Central Germany, with the consecutive Tiefstich phases of Altmark Group (ca. 3500-3350 BC), Walternienburg (ca. 3350-3100 BC), and Bernburg (ca. 3100-2650 BC, possibly commencing somewhat earlier). While the initial spread concerned HG territory, Walternienburg marks the first expansion into a traditional EEF settlement area, around the Elbe-Saale confluence. The city of Bernburg is already deep in EEF territory, and by around 3100 BC, the Tiefstich sphere reached from Stettin in the NW to the south-eastern foothills of the Harz. This expansion is believed to have caused the transformation of the Baalberge Culture aka early TRB S into Salzmünde (late TRB S) [Ironically, the village of Baalberge has a few years ago been incorporated into neighbouring Bernburg]. The eponymous Salzmünde enclosure was around 3100 BC burnt down and destroyed by a GAC – Bernburg coalition,  Afterwards, in the nearby Halle-Dölauer Heide enclosure, Bernburg people supplanted their stone cists over and across a Salzmünde graveyard, in obvious disregard of the latter.

Lüdelsen 3 megalithic grave (Altmark),
36th cBC (Source Wikimedia)

Hence, the expansionist and intrusive character of Bernburg, and also of preceding Walternienburg, is archeologically well attested.  However, little is known about their Tiefstich brethren west of the Harz. Were they also intrusive, or was it rather ideas and pots than people, which spread out southwestwards across the Lower Elbe ? If it was the former, to which extent did they absorb local HGs on their way (Neolithisation of northern Lower Saxony and the NE Netherlands is generally attributed to the TRB-Tiefstich expansion) ? Last but not least – is the full genetic story of TRB N/W already told by the Gökhem samples ? After all, these samples (slightly WHG-shifted EEF) stem from an area far outside the Erteboelle core, and as such prone to be colonised by incoming farmers without much local resistance. The fact that Gökhem has supplied the earliest Swedish megalithic burials, long before they appear in Scania, is further stressing the exceptionality of the location.

These are the questions I have in mind when speaking about an important geographical, chronological and cultural gap that needs to be filled aDNA-wise. It is ultimately about the Neolithisation of the Dutch-German plain (plus, by extension, Pomerania and the SE Baltics), and the phenomenon of megalithic burials, the most impressive monuments of the region prior to the erection of high medieval brick gothic cathedrals.

The Sorsum grave

Leine valley and Hildesheimer Wald from the SW (Source Wikimedia)

The Sorsum grave lies on the southern TRB periphery (even outside of it if you trust Bakker’s map), at the eastern side of the “Hildesheimer Wald”, a low mountain range that separates the Leine and Innerste valleys, and marks the transition from the North German plain to the Middle German low mountains. As typical for this transition belt, the surrounding valleys and the onset of the North German Plain posess highly fertile loess soils, so the Sorsum area – unsurprisingly – housed an LBK settlement chamber.

The Sorsum chamber grave
Source: Schoknecht e.a. 2016

The grave itself is not a standard megalithic burial – in fact, it isn’t megalithic at all. Instead, it consists of a 15 m long, some 2,1 – 2,4 m wide and 1,2 m deep chamber carved out of the solid rock, presumably covered by logs and overtowered with soil. The layout resembles the French “allées couvertes” and their German pendant, the “gallery graves” for which the Wartberg Culture further to the SW is famous (though there are also a handful of TRB gallery graves between the northern Harz foothills and the Saale).  Moreover, just 2 km to the west, on the mountain top, lies the “Beusterburg”, with some 20 ha size the largest Michelsberg Culture causewayed enclosure in the Lower Saxony mountains. Close spatial assosiation of Michelsberg enclosures and subsequent Wartberg gallery graves is well attested, e.g. from Calden n. Kassel.  This all gives reason to suspect a Wartberg Culture origin of the Sorsum grave. However, the 100 – 150 skeletons recovered from the Sorsum grave (often disarticulated, skulls piled up in pyramids etc.) were exclusively accompanied by Tiefstich and Bernburg pottery. Nevertheless, if there was a location where an EEF-rooted, middle-Germany MN population acculturated to  “nordic” TRB-Tiefstich influence, it should have been  Sorsum.

Weser cliff near Steinmühle (Wikimedia)

Finally, it is worth noting that the Sorsum grave / Beusterburg ensemble occupies a strategic location. The Leine valley offers the best southward transit through the low mountain ranges towards Franconia and ultimately the Adriatic Sea. It is wider and less curved than the Upper Weser valley with its many cliffs that in pre-modern times required frequent river crossings when travelling along the Weser by foot/ cart. Therefore, e.g. Roman campaigns towards N Germany preferred the Leine route, and it is also not by accident that the Hannover-Fulda (-> Frankfurt/ Munich) high-speed railway passes immediately east of the Sorsum grave.
Along the Innerste and its tributaries, the depression SW of the Harz and ultimately Thuringia and Bohemia can be reached conveniently. Northwestwards, this axis continues from Sorsum via the Weser crossing at the Müsleringen MK enclosure to the lower Ems and East/West Frisia.

Last but not least, the northern branch of the Hellweg (Duisburg – Soest -Hameln – Magdeburg) passes by Sorsum. The Hellweg forms part of what is arguably Europe’s most important West-East connection, namely Paris – Berlin – Kaliningrad – Vilnius -Moscow, the German part of which is designated as B 1 (federal road #1).  Raetzel-Fabian (2002) argues quite convincingly for the Hellweg’s existence already during the MN, as route of the Michelsberg expansion into the northern Harz foreland. Subsequently, the Hellweg served for exchange between TRB-Tiefstich and the Wartberg culture that is, e.g., demonstrated by frequent finds of axes made from “Widaer Schiefer” in Wartberg Culture gallery graves (e.g. Calden, Züschen). “Widaer Schiefer” occurs exclusively in the Harz, processing sites are evidenced from Halberstadt-Spiegelsberge (“Sonntagsfeld”) and the Jätchenberg n. Quedlinburg (Bernburg enclosure). J. Müller (2000)  uses the (non-)distribution of “Wiedaer Schiefer” axes  as one of several indications of a clear-cut “border phenomenon” between the Tiefstich (Walternienburg-Bernburg) sphere and the Salzmünde culture to its southeast. Apparently, such a clear-cut cultural border didn’t exist between TRB-Tiefstich and Wartberg. Frequent occurence of Rijkholt and Spiennes flint in the Hellweg zone indicate the westward continuation of this axis already during the MN, while occasional amber finds point to some degree of interaction with the Kaliningrad area.

In summary, Sorsum, or better the adjacent Beusterburg, is likely to have been a major trade hub that could have attracted immigrants from all directions. This means that the single sample presented below should not be over-interpreted, he may well represent an outlier. More sampling will be required to make final conclusions about TRB-Tiefstich.

The TRB-Tiefstich sample

The Krause-Kyora paper hardly discusses the Sorsum TRB-Tiefstich sample. They only state that it was a male, from whom they extracted and mapped 9,856,001 human genomic reads (1.2 fold coverage). Data has been uploaded to the European Nucleotide Archive, accession No. PRJEB24921. Maybe someone can retrieve it for further analysis (uniparental markers, f-stats etc.).

However, their Supplementary Information includes a PCA in which the Sorsum sample together with previously published ancient populations is projected on 27 modern day West Eurasian populations (not shown). This PCA, with some amendments made by myself, is displayed below. In general, I don’t have much trust in PCA analysis, as it reduces genetic complexity to just two dimensions. Even more problematic is aDNA projection on modern populations, so the PCA dimensions are determined by modern rather than aDNA diversity. Last but not least, there is no information available how much of the genetic diversity is explained by PC1 and PC2. All in all, anything but an ideal source for analysis, but the only one available so far.

The first thing that can be noted is that the Sorsum sample plots quite a bit apart from the Swedish (Gökhem) TRB samples. This indicates that Gökhem, as I have reflected about above, probably only represents a part of the genetic diversity within the TRB-Tiefstich sphere [Baalberge (early TRB S),  unfortunately not included in the PCA, was, IIRC, closer to EEF than TRB Gökhem, adding a further element of genetic differentiation inside the TRB horizon].

More specifically, TRB Sorsum appears to incorporate a substantial portion of HG ancestry, possibly in the 40-50% range when related to Anatolian farmers. In this sense, he is reminiscent of the Blätterhöhle MN samples (ca. 3600 BC, late Michelsberg). In fact, considering the Bätterhöhle’s location not too far from the Hellweg, migration from there to Sorsum is well imaginable. The find of a NW Italian Jadeite axe 2km north of the Bätterhöhle indicates connection to MN  long-distance exchange networks.
However, one may equally think of  a Blätterhöhle-like scenario, i.e. EEF from the low mountain ranges and the adjacent loess belt mixing with WHG from the North German plain, having occured along the whole northern loess border, including locally in Sorsum. Additional aDNA from comparable sites, e.g. the Belgian interaction zone between Michelsberg and Swifterband, or the Hellweg between Sorsum and Magdeburg, could provide further clarification whether Blätterhöhle is singular, or marks a general pattern of the 4th millennium BC.

Thirdly, the Sorsum sample in the PCA seems to display a slight pull towards SHG/ EHG, or maybe even the Steppe. Whether such influence is real and not just a projection artefact, and – if real – which admixture components were in play, of course requires further analysis of the whole genome. Considering that Bernburg posessed domesticated Botai horses, and there is some archeological indication of Bernburg – late Maykop contact, I wouldn’t in principle rule out some (individual) Steppe immigration into Central Europe already during the second half of the 4th millennium. More plausible, however, is a sizeable SHG element picked up in the SW Baltic Sea area, from where the Tiefstich expansion originated (and from where we unfortunately don’t have any aDNA, neither Erteboelle nor TRB, so far).

Whichever influence(s) differentiated TRB-Sorsum from TRB-Gökhem, and even more from TRB-Quedlinburg (Baalberge/ early TRB S; 110 km and  ~400 years away) – he clearly represents a substantial intrusive element into this traditional EEF settlement chamber.

Another interesting observation is that Unetice in the PCA plots about half-way between Yamnaya and the Sorsum sample. CWC, in comparison, drifts towards SHG (or Narva ?). Of course, this observation requires confirmation by formal testing of the full genome. If it stands such test, this would indicate that the Sorsum sample isn’t an outlier, but quite representative of the population in the Dutch-German plain and adjacent areas, which admixed with incoming Yamnaya-like populations during the LN to ultimately form Unetice, and to a lesser extent also participated in the formation of CWC. This population may plausibly be labelled as TRB-Tiefstich, including Bernburg as late offspring, maybe plus a Blätterhöhle-like element in Wartberg.

Let me finally remark that according to the study’s figure supplements 8/9, TRB-Sorsum is projected in-between modern non-Basque Spanyards, and a cluster of modern Tuscans, Albanians and Bulgarians. Of course, this shouldn’t be over-interpreted (“Gothic motive“, etc.), but is probably owed to the fact that the Sorsum sample, as most South European populations, comprises very little Steppe ancestry. Nevertheless, it is noteworthy that – in contrast to EEF, Yamnaya, and also most CWC samples – TRB-Sorsum is projected inside the genetic variation of modern Europeans.

Let’s take that final remark a bit further: TRB is generally understood to have emerged from the interaction of the Michelsberg Culture with the Epi-Lengyel horizon (Gatersleben in Middle Elbe-Saale, Jordanow in Bohemia and Silesia, late Lengyel in Kujawy). Re-phrasing an intentionally simplifying remark by J. Müller: If you combine flat-based epi-Lengyel pottery with Michelsberg tulip beakers, you get Funnelbeakers. While the Michelsberg expansion was long believed to have stopped somewhere west of the former Iron Curtain, early MK presence (ca. 4000 BC) in Thuringia, Saxony and Bohemia, plus tulip-beaker inspired forms in Silesia, are now archeologically well attested; recent finds suggest that MK may also have reached the lower Oder.  This yields a quite large zone of  epi-Lengyel – MK interaction that stretches between Prague and Stettin, and possibly extends to W. Sweden.

Michelsberg is believed to have emerged from the Paris basin, w. backward linkage to Brittany, incorporating epi-Cardial and pastoralist HG (La Hoguette) elements. Lengyel had a Danubian origin and built on the Vinca metalurgical tradition. The Sorsum sample additionally suggests a sizeable WHG/SHG component in the TRB Tiefstichkeramik sphere that isn’t fully represented by the Gökhem samples, and most likely represents Swifterband-Erteboelle adstrate. In summary, TRB-Tiefstichkeramik may represent the fusion of both of the EEF streams (Mediterranean/Cardial, and Danubian/LBK, including their differing HG substrate), with North European post-Kongemose HG traditions.  Even though it took to the LN to complete the mix with a fourth element, namely Steppe-like ancestry from Eastern Europe, the three a/m components should already have combined sufficient ancestry sources to yield a “modern European” – like result. And, there is hardly a better place imaginable for such a fusion to materialise than Hildesheim (-Sorsum), at the junction of the European main East – West (Moscow – Paris) and North – South (Norway – Sicily,  modern E 45) communication axes.

11 thoughts on “A first, and intriguing glimpse at TRB West Group aDNA

  1. Frank, those are some amazing maps. Well done
    Can you perhaps sketch out a bit how the succession of TRB phases proceeded ?
    Is Baalberg an early phase of TRB (if so which subgroup), or a pre-TRB culture ?

  2. It’s a shame we don’t have that sample to look at it in more detail. Now I was looking at the Blätterhöhle samples to see if they had some “eastern” shift:

    Blatterhole_MN:I1563
    Barcin_N 55%
    WHG 22.2%
    Koros_HG 17.5%
    Baltic_HG 5.3%
    Latvia_HG 0%
    EHG 0%
    Yamnaya_Samara 0%
    CHG 0%

    Distance 3.5581%

    Blatterhole_MN:I1593
    Barcin_N 52.6%
    WHG 47.4%
    Koros_HG 0%
    Latvia_HG 0%
    EHG 0%
    Yamnaya_Samara 0%
    CHG 0%
    Baltic_HG 0%

    Distance 3.3585%

    Nothing striking apart from the already known high HG admixture. But look at the sample labelled as HG:

    Blatterhole_HG
    WHG 47.5%
    Baltic_HG 31.3%
    Barcin_N 15.2%
    Yamnaya_Samara 3.1%
    Koros_HG 1.9%
    Latvia_HG 1%
    EHG 0%
    CHG 0%

    Distance 2.1228%

    Blatterhole_HG
    WHG 43.5%
    Baltic_HG 31.7%
    Globular_Amphora 21.1%
    Yamnaya_Samara 2.9%
    Latvia_HG 0.8%
    Koros_HG 0%
    EHG 0%
    Barcin_N 0%
    CHG 0%

    Distance 1.8771%

    This one does look interesting. I’ll have to check the exact date of the sample.

    Looking again at the 2 MN samples, but adding other MN sources:

    Blatterhole_MN:I1563
    France_MLN 33.2%
    Baltic_HG 26.2%
    Globular_Amphora 20.1%
    Germany_MN 18.9%
    Koros_HG 1.1%
    Yamnaya_Samara 0.5%
    WHG 0%
    Latvia_HG 0%
    EHG 0%
    Barcin_N 0%
    CHG 0%
    LBKT_MN 0%
    Iberia_MN 0%

    Distance 2.1064%

    Blatterhole_MN:I1593
    France_MLN 30.8%
    WHG 25.3%
    Iberia_MN 22%
    Globular_Amphora 10.9%
    Baltic_HG 7.4%
    LBKT_MN 2.6%
    Germany_MN 1%
    Koros_HG 0%
    Latvia_HG 0%
    EHG 0%
    Yamnaya_Samara 0%
    Barcin_N 0%
    CHG 0%

    Distance 1.9363%

    Both get a good amount of France_MN (and one also Iberia_MN), but then also some Globular Amphora and Baltic_HG. I guess I’ll need to run a more comprehensive set of tests to see what’s the consistent pattern and best fits.

  3. Those Blatterhohle samples all date to same period, 4000-3000 BC. I’d guess Davidski simply labelled one as “HG” in his G25 by virtue of his high HG ancestry, not that it dates earlier than the other 2 or is actually from the Mesolithic per se.
    Not much at all was written about their context in Lipson et al (although we are thankful for such samples), and I dug up a brief report on Blatterhohle recent excavation. They were cave burials, with rather undiagnostic pottery, and the middle Neolithic time frame was concluded by C14 dating. So I imagine this is research still in progress.
    I’m not convinced what much else we can tease out in terms of intra-European middle-Neolithic relations, certainly using GAC, France_MLN poses the problem of contemporeity (in fact the GAC individuals post-date them). Other approaches might entail by looking at Y -DNA patterns (which I might get to one day) or some clever D-Stats.
    Compared to classic earlier ‘WHGs’ , the Blatterhohle outlier does have some eastern shift, maybe from Iron Gates ? Pre-Neolithci hunter-gatherer networks can probably explain it.

    But what these do show is that near-pure hunter-gatherer individuals existed into the 4th millenium , unadmixed, in northern Europe, which account for the high WHG-admixed samples which subsequently emerge, such as also the individual at Sorsum. I don’t think any European archaeologist would be too surprised by such finds, but it’s great to be confirmed.

    Some distant models:

    Population EHG CHG WHG Balkans_N Iron_Gates_HG Ukraine_Mesolithic Baltic_HG:Spiginas4 Distance

    Germany_MN:I0172 0 0 16.8 83.2 0 0 0
    0.03683
    Germany_MN:I0560 0 0 19.9 80.1 0 0 0
    0.027971
    Blatterhole_HG:I1565 0 0 44.9 8.8 33.6 2.4 3.9
    0.018861
    Blatterhole_MN:I1563 0 0 21 58.1 0 0 20.9
    0.031636
    Blatterhole_MN:I1593 0 0 46.1 53.8 0 0 0
    0.02951
    France_MLN:I4303 0 0 20 80 0 0 0
    0.051504
    France_MLN:I4304 0 0 20.5 79.5 0 0 0
    0.044375
    France_MLN:I4305 0 0 19.2 80.8 0 0 0
    0.043358

  4. A. Baalberge:
    First note the plural; Baalberge . Originally, there were several burial mounds which lent their name to the village (Baal stands for pre-/ non Christian, -berg = “hill”). Only one mound, however, has remained to date.

    Baalberge is early TRB S Group (with Bakker’s map being somewhat inprecise). Around the turn of the Millennium, J. Müller dated it in Middle-Elbe Saale (MES) to ca. 3800-3400 BC. In the meantime, additional finds give reason for revising that chronology. The Hundisburg enclosure n. of Quedlinburg, e.g. has yielded mostly Baaalberge sherds (plus an occasional MK tulip beaker), and has been dated to 4200 – 3800 BC. Bohemians are more clever in this respect – they have TRB around 4000 BC commencing with a “Michelsberg-Baalberge Phase”. Baalberge proper commences around 3700 BC there, that is also when it starts to show in Moravia. NE Brandenburg & W. Pomerania (TRB N / TRB E acc, to Bakker) has also yielded early Baalberge finds (39th cBC). It is unclear to me how long it lasted there – ultimately it was replaced by TRB-Tiefstich, but possibly not before ca. 3300 BC.

    In MES, some authors define additional TRB subgroups, namely Schöningen-Schiepzig (northern Harz foreland, possibly commencing as early as 4200 BC) and Hutberg (Middle Saale/ Thuringia, Baalberge-Salzmünde transition). Their specificness has remained unclear to me. From some papers, it seems that, in addition to TRB, they also contain some Michelsberg (MK II-V), Epi-Lengyel (Gatersleben/Jordanow) and possibly (proposed by Raetzel-Fabian) Baden-Boleraz elements.
    IMO, instead of a homogenous “culture”, however, we may rather be dealing with multi-cultural setups at major proto-urban communication centers. Schöningen, e.g., an ancient “salt town”, in my map is the big junction 50 km N of Quedlinburg. Salzmünde-Schiepzig is the location of the Salzmünde enclosure – to avoid confusion, the early settlement phase here has been named as Schiepzig group, the later one as “Salzmünde”. The Wallendorf-Hutberg enclosure lies 20 km W of Leipzig. The second largest “Hutberg” findspot is the Halle-Dölauer Heide enclosure, 5 km SE from Salzmünde (strongly fortified, 25 ha),

    The late phase of TRB S is Salzmünde, believed to have started around 3400 BC in MES, 3300 BC in Bohemia. There is little, if at all, Salzmünde in Moravia, where TRB fell victim to Baden-Boleraz. From the north, it got crushed by the Tiefstich expansion.

    B. Blaetterhoehle (not Blatterhole, lit. “leaves cave”):
    Indeed, the cultural assignment to Late Michelsberg / early Wartberg has essentially been made based on temporal and geographical considerations. However, the finds aren’t as “undiagnostic” as it may seem at first glance:

    1. Culturally, MK use of caves is quite well attested, but didn’t receive much attention before the Blätterhöhle finds. Cases have, e.g. been reported from the Harz (“Baumannhöhle”), the Swabian Alps, and Upper Franconia. However, from what I could find, only the “Jungfernhöhle” (Bamberg county) has provided human remains (ca. 35 skulls – yet undated), Across the Rhine, cave burials are quite common in SOM, especially in the Ardennes (BE/LUX).

    2. Spatially, Blätterhöhle falls well within the MK/ Wartberg area. A MK hilltop settlement is attested some 8 km further east (Burgberg Oestrich), and the nearby terraces of the Ruhr and Lenne rivers have yielded various MK stray finds. Blätterhöhle lies at the outskirts of Hagen, a city that owes its existence to control over northward transport of the Siegerland’s iron resources along the Lenne towards the Ruhr area industrial belt. Iron smelting in the Siegerland, of course, “only “dates back to the Iron Age. but use of Siegerland iron ore as color pigment, prepared as (lip?-)stick, is already archeologically attested from the MN onwards (Netphen, various Rössen and MK finds). The Blätterhöhle surrounds have yielded evidence of long-distance connections, e.g. a Piemontese (Monte Viso) Jadeite axe, and tools made from Belgian flint.

    Against this background, there are two questions to answer:
    1. What background had the EEF-like portion of the Blätterhöhle admix? More specifically, does it rather look Western/ Cardial, thus confirming the suspicion that MK (possibly also preceding Rössen) represents demic expansion out of Eastern France, or Danubian /LBK like ?
    Your stats, Alberto, seem to point at the former. May I suggest you, Rob, to incorporate post-Cardial (e.g. Iberia_MN) in your distant models?
    2. What about the admixing HGs? If it was a surviving HG population from the Westphalian uplands, it should essentially resemble WHG (Loschbour ca. 250 km SSW of Blätterhöhle). If, OTOH, we are dealing with Tiefstich-like expansion from the SW Baltic Sea, we might expect substantial Baltic_HG and /or SHG elements (the latter is yet missing from both your, Alberto and Rob, analyses).
    And, in fact, the Blätterhöhle_HG and at least one of the Blätterhöhle_MN samples appear to contain a “Baltic” signal.

    C. Esperstedt_MN (I 0172)
    I had another look at the sample’s description in Haak e.a. (2015). What struck me there was that it came out as ~40% WHG (Fig. 3). They had problems with the cultural assignment: The archeological context was Bernburg-like, but AMS dating (3360-3086 BC) placed it into the Salzmünde period.
    A possibility to envisage is that the Bernburg chronology requires upwards revision. A German presentation abstract describes the 32nd cBC Tiefstich-Salzmünde boundary as running along the line Erfurt-Potsdam-Stettin, which might (just) place Esperstedt within the Tiefstich sphere. Müller’s 2000 analysis (linked in my post) suggests the same.
    You, Rob, however, analyse Esperstedt_MN as 17% WHG – 83 % Balkans_N, which substantially differs from Haak e.a. Would you mind running another check on that sample, possibly also including a “Western” EEF source (MK proxy), as well as SHG?

  5. Thanks for putting that up Alberto

    Frank, based on this it would seem the majority of ancestry of French MN and large parts of Michelsberg would be from Iberia. Indeed, LBK seems to have collapsed, with hiatus in areas like Alsace, the middle Rhine, etc, and perhaps the new stream was mostly Iberia/ Cardial ? Yet LBK shows a continuity in the Paris basin, by way of RRBP and BVSG..
    But then again, G25 could be just preferring it due to the higher WHG in Iberia MN c.f. LBK.
    I think in the works are genome wide data from France which will help clarify this question.

  6. Rob:
    There was a study on RRBP that, based on uniparental markers, suggested RRBP having formed by merger of LBK with Cardial-influenced hunter-pastoralists such as La Hoguette.
    OTOH, archeobotany suggests that some Danubian EEF reached NE Iberia, where they, together with local HGs, may have contributed to the transformation from Iberia_EN to Iberia_MN [Alberto, that is more your turf, can you confirm or decline?]
    As such, one does not necessarily have to assume a direct Iberian relation to Central Europe. Instead, Iberia_MN may reflect a more widespread merger of Danubian, Cardial, post-Magdalenian and post-Epigravettien elements that occured West of the Rhine.
    According to Lipson (2017), Iberia_MN includes significant La Brana admixture. You could do a test run that includes La Brana, but excludes La Brana-loaded Iberia_EN/MN, to see if it shows up in TRB samples. Alternatively, Iberia_EN may be used instead of Iberia_MN to test for a “Cardial” signal.

    LBK did not only collapse along the Rhine. Brotherton (2013) showed for MES that LBK mtDNA was almost completely replaced during the MN, starting already with Rössen. A similar finding was, IIRC, reported for Hungary. Replacement of LBK yDNA with yhg I2a by TRB (or already MK?) has anyway become obvious by now. The only area where LBK, in its final, Bohemia-originating stage of Stichbandkeramik (SBK, “stroke-ornamented pottery” or “pass linear pottery”) seems to have survived is Bavaria and (Western) Bohemia. Substantial SBK substrate is the preferred explanation for that region’s specific MN cultures such as Altheim or Cham.

    Studies (e.g., Ostritz 2003 http://www.jungsteinsite.uni-kiel.de/pdf/2003_ostritz.pdf ) indicate substantial decrease in MES settlement density already during the LBK-SBK transition. Later newcomers, especially Rössen, didn’t “take over” LBK settlements but mostly occupied vacated areas . This doesn’t rule out occasional violent conflict, but such conflict is unlikely to have been the main reason for LBK virtually “dying out”.

    I rather suspect epidemics, especially the flu. Influenza viruses are best adapted to surviving in cooler water (< 17° C) with low salinity (as, e.g., the Baltic Sea), major hosts are aquatic birds and sea mammals. [A case of deer-to-human flu transmission in the 1990s has also been documented for N. Germany]. Seal-hunting Erteboelle people should have acquired a decent degree of immunity, possibly comparable to modern North Europeans. For ANF, however, originating from a much warmer and dryer climate zone, exposure and acquired immunity should have been much lower, especially as concerns seal flu viruses (c.f. https://www.webmd.com/a-to-z-guides/news/20120731/seal-flu-next-pandemic-threat). This applies even more to inland-settling Danubian EEF. Close contact to pigs, a well documented secondary Influenza host, should have added further to the infection risk.

  7. @ Frank

    1) “”LBK did not only collapse along the Rhine. Brotherton (2013) showed for MES that LBK mtDNA was almost completely replaced during the MN, starting already with Rössen. A similar finding was, IIRC, reported for Hungary. Replacement of LBK yDNA with yhg I2a by TRB (or already MK?) has anyway become obvious by now. The only area where LBK, in its final, Bohemia-originating stage of Stichbandkeramik (SBK, “stroke-ornamented pottery” or “pass linear pottery”) seems to have survived is Bavaria and (Western) Bohemia. Substantial SBK substrate is the preferred explanation for that region’s specific MN cultures such as Altheim or Cham.””

    Yes the SBK does seem typologically continuous, or evolved from, LBK. There’s not much G-W data from that group yet.
    About TRB, as we know, there is quite some gap between TRB (c.4000->) and LBK. So some intervening cultures like Epi-Rossen , Gatersleben would be instructive to understand how the shift happened.
    I think it’ll take more detailed data sets than just mtDNA to see if Brotherton’s conclusions about a discontinuity are upheld, as it might be a matter of hunter-gatherer introgression rather than abrupt shift. Don’t archaeologists suggest Rossen evoloved typologically from LBK ?

    About Hungary – the transition post Early Neolithic is regionally variable. I don’t think we can invoke one single meta-narrative .

    About Iberia, a very interesting abstract listed for the upcoming Jena meeting .
    The late Mesolithic shift in Iberia toward central European Mesolithics (?””Castelonovization”), presumably away from the Magdalenian-inclined La Brana types would would only further complexify the issue of post-Cardial ancestry throughout Europe.

    The genetic history of the Iberian Peninsula over the last 8000 years.

    ”The Iberian Peninsula, lying on the southwestern corner of Europe, provides an excellent opportunity to assess the final impact of population movements entering the continent from the east and to study prehistoric and historic connections with North Africa. Previous studies have addressed the population history of Iberia using ancient genomes, but the final steps leading to the formation of the modern Iberian gene pool during the last 4000 years remain largely unexplored. Here we report genomewide data from 153 ancient individuals from Iberia, more than doubling the number of available genomes from this region and providing the most comprehensive genetic transect of any region in the world during the last 8000 years. We find that Mesolithic hunter-gatherers dated to the last centuries before the arrival of farmers showed an increased genetic affinity to central European hunter-gatherers, as compared to earlier individuals. During the third millennium BCE, Iberia received newcomers from south and north. The presence of one individual with a North African origin in central Iberia demonstrates early sporadic contacts across the strait of Gibraltar. Beginning ~2500 BCE, the arrival of individuals with steppe-related ancestry had a rapid and widespread genetic impact, with Bronze Age populations deriving ~40% of their autosomal ancestry and 100% of their Y-chromosomes from these migrants. During the later Iron Age, the first genome-wide data from ancient non-Indo-European speakers showed that they were similar to contemporaneous Indo-European speakers and derived most of their ancestry from the earlier Bronze Age substratum. With the exception of Basques, who remain broadly similar to Iron Age populations, during the last 2500 years Iberian populations were affected by additional gene-flow from the Central/Eastern Mediterranean region, probably associated to the Roman conquest, and from North Africa during the Moorish conquest but also in earlier periods, probably related to the Phoenician-Punic colonization of Southern Iberia.”

  8. Rob “Don’t archaeologists suggest Rossen evolved typologically from LBK ?
    Well… Rössen in general displays typical EEF features: Settlement of loess areas <350 m a.s.l., flat grave inhumation (albeit stretched on the back in proto-Rössen Hinkelstein-Großgartach, in contrast to LBK positioning the deceased flexed on the side), longhouses (in fact, now "giant houses" of up to 65m length). One may even argue that the LBK-SBK shift was more pronounced, with the latter switching from rectangular to trapezoid longhouses, and introducing astronomical rondels ("henges", e.g., the Goseck circle) – traditions continued by Rössen.

    However, while SBK continued LBK settlements, Rössen systematically avoided any locations on which LBK/SBK had settled before (or were still settling). This even concerns LBK/ SBK enclosed settlements, typically major communication centers in strategic locations. Instead, they set up their own enclosures nearby. An example is North Bochum, with a Rössen "giant house" on the "Hillerberg", in 2.5 km distance to the large, enclosed Bochum-Bergen LBK settlement (which btw delivered samples of the Siegerland iron oxide pigment sticks that I have discussed above) . Such scruples were unknown to subsequent cultures – Gatersleben and Baalberge, e.g., continued to use the enclosure and cemetary of the eponymous village of Rössen, Bernburg "recycled" the Eilsleben LBK enclosure, Halberstadt-Sonntagsfeld (Spiegelsberge) displays long-term post-Rössen settlement continuity, etc. So, in some way, Rössen people must have felt "alien" to LBK/SBK, in spite of sharing (or adapting to?) a similar cultural background.

    Rössen (ca. 48th – 45th cBC) goes back to the Hinkelstein Group that appears around 5000 BC, still contemporary with later LBK, on the northern Upper Rhine. Subsequent Großgartach (GG, from 4900 BC) replaces LBK along all of the Rhine, including the Rhineland, Alscace, and the Neckar valley, in Swabia and Franconia, while SBK succeeds LBK in Bohemia, SE Bavaria, MES and Poland, Early Lengyel in Transdanubia and Moravia, and (Pre-) CT in Moldova and Ukraine. As SBK, GG-Rössen builds astronomical rondels/henges, e.g. Ippesheim (GG), Bochum-Harpen (Rössen), pointing at some shared ideology that possibly emerged from Hungarian Early Lengyel (https://www.researchgate.net/publication/278658643_Neolithic_Circular_Ditch_Systems_Rondels_in_Central_Europe ). The shift from rectangular to trapezoid houses is also common to both SBK and GG – Rössen (no Hinkelstein houses have been found yet), with the direction of innovation transfer yet unclear.
    Rössen expanded even further than GG- aside from entering MES (Rössen is a suburb of Leuna, at the western bank of the Saale) where it co-habitated with SBK, it also spread into Luxemburg and Dutch Limburg. Rössen pottery was furthermore found in the Lower Saxony plain (e.g. Hüde/ Dümmer), Hamburg, on the Lower Oder, and in Dabki on the E. Pomeranian coast. [That last site, a pastoralist-HG settlement with syncretic mix of Erteboelle, Narva, SBK and Rössen pottery, arguably also some Bodrogkeresztúr imports, is a key place when it comes to neolithisation of the Central Baltic Sea area, and formation of the TRB phenomenon; it has delivered very early (~4100 cal.BC) TRB pottery]. Moreover, Rössen-style decoration occurred in the Franche Comte, the Paris Basin, SE Bavaria, and Poland, where it sometimes lived forth after 4500 BC, e.g. in the primarily Lengyel-influenced Bresc-Kuyavsky Culture (c.f. L. Czerniak “After the LBK”, 2010).

    As such, the intrusive character of Rössen in MES is archeologically clear. Unclear, however, is yet whether Hinkelstein on the Northern Upper Rhine displays the original starting point, or in itself reflects a movement that commenced elsewhere. Hinkelstein burials in extended supine position are reminiscent of Mesolithic traditions (the same, btw, applies to TRB flat graves, also typically w. extended supine burials). Since the Northern Upper Rhine was traditional EEF territory, the custom looks likely to have originated elsewhere. The culture itself derives its name from a Menhir (“Hinkelstein” in German) that marked the first graveyard excavated near Worms.. There are several Menhirs in Germany, but most appear to stem from later periods (e.g. Langeneichstädt, next to a Salzmünde/Bernburg gallery grave, or the “Steinerne Jungfrau” in the Salzmünde enclosure of Halle-Dölauer Heide). Contemporary Menhirs are best known from Brittany.

    I think it’ll take more detailed data sets than just mtDNA to see if Brotherton’s conclusions about a discontinuity are upheld, as it might be a matter of hunter-gatherer introgression rather than abrupt shift
    If HG introgression played a role, it wasn’t local one that, for all we know, should have been mtDNA U5. In Brotherton’s dataset, LBK had less than 20% mtDNA H, going up to 36% in Rössen (though the Rössen subtypes are rather undiagnostic of origin – H1, H5b, H16, H89). For Salzmünde (~30% H), they report two cases of H3 that, for all we know, was native to the Gulf of Biscay area. However, those women may also have arrived with Michelsberg. Modern Central Europeans have a frequency of some 40% mtDNA, and that is neither due to Central/ North European HGs, nor to Steppe/ CW, where mtDNA H played only a minor role. The main factor, as per Brotherton, was BB at ~50% mtDNA H, nearly half of which consisted of “Iberian” H3 and H5a (apparently steppe-related BB men had a crush on Iberian women …)

    However, I agree that Rössen deserves more in-depth analysis. Lipson (2017) includes the first and so far only full Rössen genome, which, unfortunately, has received even less attention so far than Blätterhöhle.

  9. Frank,
    In ”Hinkelstein- The cultural picture..”, Fischer points out that their pottery is derived from Stroked-Ornamented culture, and as you say, assoc. with a shift to extended inhumation.
    In the Alsace culture project, Denaire et al point to a 150-300 year hiatus between the end of the latest LBK phase and the beginning of Hinkelstein.
    As you suggest, the Rondel tradition points to radiation of a cultural idea from the Lengyel core zone in Austria & transDanubia.
    Lastly, the elevated mtDNA H need not come from Iberia but could come from the Carpathian region (a/p Harvella et al).
    Putting this together, it could be that the early LBK was replaced from a second wave from the middle Danube region to the Rhine, starting c. 49-4700 BC.

    Further questions/ issues:

    – as Pasztor asks in your linked article, how do Megaliths e.g. in Britain link to this, given their chronological disparity (ie. are much later).
    Mueller and Furholt previously suggested that as the Rondel idea reached France, it began to transform, from just rituals to burial., the Passy type ditches with burials, before transforming further still into Megalithic ideas in concert with local Mesolithic substratum, at the end of the 5th M in Atlantic France.

    – how do Michelsberg ditches relate to Rondels and ditched enclosures of the Lengyel-circle type ?

    – at the other end of Europe, it has been pointed out (Manzura) that the first true monumental kurgans in the westernmost steppe, could be inspired from the astronomical ideas of Rondels, their ditch-palisade systems, etc.

  10. Rob:
    In ”Hinkelstein- The cultural picture..”, Fischer points out that their pottery is derived from Stroked-Ornamented culture
    And in his classic paper “Die Stichbandkeramik im Saalegebiet” (1974), D. Kaufmann demonstrates that SBK trapezoid houses only appear in what he calls SBK Ib, and become dominating in SBK II, which he explains with significant Hinkelstein / early Rössen influence. SBK II also sees appearance of stretched supine burials.
    While differentiating culturally, Rhine and Elbe were still connected via various systems of cultural and material exchange. Most significant in this respect is Arnhofen Amphibolite from the Regensburg area,that was traded along the so-called “flint route” to Pilsen and Prague, and also found in GG burials at Trebur. [Unsurprisingly, the SE Bavarian MN presents itself as a fusion of SBK and Hinkelstein-GG influences]
    Kaufmann also points out that some proto-forms of stroke-ornamented pottery developed in the late LBK around the Lower Main, from where they may independently have spread to Bohemia and MES (and of course also to Hinkelstein just across the Rhine).

    J.P. Farrugia “Une crise majeure de la civilisation du Néolithique Danubien” even goes a step further by postulating a “tabooing” of curvilinear LBK decoration around 5100 BC that according to him originated in E. Hungary and quickly spread across most of the LBK oikomene. This “tabooing” forms part of the evidence he tries to construct for a LBK “civil war” around 5100 BC. (As interesting as this “civil war” theory is, I propose to not discuss it further here). In any case, such “tabooing” would automatically lead to tri- or rectangular decoration patterns, which of course are more suitable for using (bone-made) comb-like decoration tools as those known from SBK. Farrugia also points at early occurence of stroke-ornamentation in the Aisne valley, and reminds of such decoration being typical for La Hoguette.

    In short – this seems to be a classical case of pre-aDNA / pre-AMS dating archeological debate, in which everybody is free to extrapolate his local excavation experience to the larger Central European context. R. Gleser (2010, http://www.academia.edu/3397719/Zeitskalen_stilistische_Tendenzen_und_Regionalit%C3%A4t_des_5._Jahrtausends_in_den_Altsiedellandschaften_zwischen_Mosel_und_Morava ) has on 80 pages tried to somehow sort out chronological (and implicitly also cultural) relations between the various Central European post-LBK Groups, to ultimately concede failure at least as concerns the 1st half of the 5th mBC, a/o for lack of reliable AMS dating (especially SBK), and a tendency of local researchers to construct clear-cut sequences out of patterns that may sometimes be better interpreted as poly-cultural.

    More substantial than Fischer’s paper, anyway, is Helmut Spatz: “Bäumchen und Sichel” published in the same journal. Spatz excavated the Hinkelstein-GG graveyard at Trebur, and most of Fischer’s statements go back to Spatz’ findings which he presents clearer and in more detail in the a/m publication than Fischer does. Aside from discussing parallels, and differences, between SBK and Hinkelstein – GG pottery decoration, Spatz highlights the following:

    1. “Distinct features of the economy of both the Hinkelstein (with stock–raising and hunting) and Grossgartach (agriculture) cultures can be recognised ” His arguments for a pastoralist-HG Hinkelstein background include
    (i) meat deposits, up to whole pigs and halved cows, in over 50% of Hinkelstein burials – a custom rarely known from LBK, and also GG burials, but present in Rössen and later MN cultures (GAC !);
    (ii) Strong representation of jewelry made from deer tooth and antler (again rather uncommon in LBK/ GG),
    (iii) Low degree of skeletal lesions typically associated with agricultural activities,
    (iv) Manifold presence of arrowheads (trapezoid, differing from LBK triangular arrowheads, but typical for MES Rössen and several later MN cultures) in Hinkelstein graves, while arrowheads were completely absent from surrounding GG burials;
    (v) Substantial wealth differentiation among the buried, including richer male than female graves, as typical of pastoralist societies, contrasting with the rather egalitarian GG burials.

    2. “Grossgartach [..] (was) constituted by the remaining (acculturated) Linear Pottery population”. Aside from the a/m differences, and Trebur GG burials including sickles as grave goods, he points to other locations where GG burials contained (LBK-typical) triangular arrowheads, in contrast to Hinkelstein trapezoid arrowheads.
    Schwarz, in general a proponent of a clear Hinkelstein-GG chronological sequence, admits in this context that Hinkelstein and GG may have co-existed – a view voiced by J. Müller when considering that AMS datings of both Hinkelstein and GG are quite dispersed and tend to overlap.
    If so, the question arises whether Rössen, which shares more similarities with Hinkelstein than with GG, wasn’t a direct Hinkelstein offspring. The answer, IMO, is rather to be found via aDNA than yet another comparison of pottery sherds.

    it could be that the early LBK was replaced from a second wave from Hungary to the Rhine, starting c. 49-4700 BC.
    A second wave of Hungarian pastoralist-HGs with Mesolithic burial traditions, leaving no trace of such burials along the upper Danube and the Neckar before arriving at the northern Upper Rhine valley? Come on! Hinkelstein “smells” post-La Hoguette in various respects, including pottery decoration style (albeit not forms), and subsistence mode, and La Hoguette had been present in the Hinkelstein area prior to LBK arrival.

    Let me add, that in the Olalde e.a. BB paper, Table S5, Germany_MN is shown with 2.4% La Brana admix (France_MLN 16.6%, GAC 3.6%, no La Brana in LBK, Gökhem). These figures all relate to ANF and KO1 as additional WHG source, p-values are >0.05. ANF – La Brana – Loschbaur models fail for Germany_MN and GAC.
    This suggests that sometime before ca. 3,500 BC, a Iberia_MN/ France_MLN related population entered Central Europe – either with Hinkelstein/ Rössen, or with MK. The fact that Gökhem, which in all likelyhood should reflect colonisation by a Michelsberg-related Group, doesn’t show a La Brana signal, makes me tend towards Hinkelstein/ Rössen. However, ultimately we need more aDNA, especially from Michelsberg, and a closer look at that Rössen sample from Lipson 2017 would also not do harm.

    I will seperately commment on your other issues.

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