Ancient DF27

ORIGINS OF DF27

Ancient phylogeny: R1b-M269 > R1b-L23 > R1b-L51 > R1b-P310 > R1b-L151 > R1b-P312

McDonald age estimates for R1b-P312 subclades: link

The commonly accepted view is that R1b-P312 had prehistoric origins in early Corded Ware Culture somewhere in the east of Europe on the forest steppe - perhaps coming from earlier Yamnaya-related or Sredny Stog culture that had origins even further east.  'Yama' is a Russian adjective that means 'related to pits' - the people used to bury their dead in burial mounds containing pit chambers.  The burial mounds are called howes (Old Norse haugr) here in Yorkshire, also known as barrows further south, and are still prominent in the moorland landscape.  R1b-M269 was present in Yamnaya populations.  R1b-Z2013, a Yamnaya  subclade, descends from R1b-M269. 

Chalcolithic-to-Bronze Age European R1b-P312 burials are the ones being found to have significant steppe autosomal ancestry.  Non-R1b-P312 European farmer burials at that time generally did not have steppe autosomal ancestry.  The paternally-related R1b-L51 clans moved west across Europe through Corded Ware territory bringing steppe admixture with them.  A local variant of Corded Ware Culture became the Single Grave Culture in northwestern Europe.  R1b-P312 was a major yDNA lineage of the Bell Beaker people.

Map of Corded Ware culture (above) via Wikimedia Commons.

Ancient yDNA from Central European (Czech Republic/Bohemia) burials studied in Dynamic changes in genomic and social structures in third millennium BCE central Europe, Luka Papac et al., 2021 included U106 burial PNL001 dated to around 2900 BC (2914 to 2879 cal BCE) and a L151 burial OBR003 (xU106, xP312) that was also dated to about the same time (2911 to 2875 cal BCE).  OBR003 had a plain-looking beaker, a type A-battle axe, flint dagger and a T-shaped antler plate of a type found in early Corded Ware from NE Poland, Latvia and Central Germany.  His autosomal DNA suggests an origin in the northeast of Europe too and both PNL001 and OBR003's autosomal DNA results plot closest to Yamnaya/steppe samples.  These dates indicate that R1b-U106 was probably born in around 3000 BC and R1b-L151, the ancestor of R1b-P312, probably originated c. 3100 BC.  This ancient yDNA dating is in the same region as the date arrived at for the age of DF27, 'son' of P312, (c. 3000 BC), estimated using SNP counting for early Rox2 NGS tests in 2014.  As ever, all dates have a margin of error.

The Genomic Formation of Human Populations in East Asia, burial I6222, from the Afanasievo (c. 3300 to 2500 BC) culture in Mongolia, is dated 3316-2918 calBCE  and is listed as R1b1a1a2a1a (R1b-L52).  FTDNA later ran analysis and called I6222 as R-P310*.  Ancient genomes reveal social and genetic structure of Late Neolithic Switzerland by Anja Furtwängler et al., April 2020, includes an autosomally Corded Ware-like R1b-L151* burial on the Swiss Plateau.  The burial is named Aesch25 and dated to 2864-2501 cal BC.  He appears to be a 'newcomer', the other men were haplogroup G2a, and he was buried with them as the last burial in a collective megalithic tomb known as a dolmen.  They say:

over time at an individual level, it becomes apparent that this ancestry [Yamnaya-related/steppe] was virtually absent before 2700 BCE, followed by a steep increase in parts of the population starting around 2700 BCE.

Corded Ware cultural complexity uncovered using genomic and isotopic analysis from south-eastern Poland by Ana Linderholm et al., published in Nature in April 2020, found R1b-M269 and two R1b-P310 Corded Ware burials in SE Poland.  Ages of the R1b samples in the paper range between 2500 BC - 2350 BC.  Proto-Nagyrev I7043 and I7041 from Olalde et al.'s Bell Beaker paper who are R-L151 (xP312,U106) and dated to 2500 BC - 2200 BC.  

An October 2019 study found a R1b-P310 Corded Ware child's burial, Tauber_CWC:ALT_4 , in the Tauber valley (Taubertal), Germany.  The burial dates to 2570-2458 cal BC and analysis of degraded ancient DNA reveals he was positive/derived for P310 and negative/ancestral for U106, with no read for P312 or other downstream SNPs.  The date is contemporary with the earliest U152 burial yet known (RISE563 c. 2542 BC).  U152 is a 'brother' of DF27 and is descended from their R1b-P312>ZZ11 'father'.  It seems possible therefore that P310* Tauber_CWC:ALT_4  and others could also be P312+ and positive for a downstream SNP that is difficult to pick up or has not been covered by the test (DF27).  The nearby Black Forest region coincides with a potential place of origin (analysis of oxygen isotope zone characteristics) for the Amesbury Archer (also R1b-P310*) and the Boscombe Bowman (FGC11381+/Y8397+ but listed as R1b-P310*) migrant burials of Stonehenge.  Those two important English Bell Beaker burials, both with high 'southern' autosomal DNA and no SNPs downstream of P310 present in their low resolution samples (other than FGC11381 for the Boscombe Bowman), are mentioned in more detail on the Bronze Age DF27 page.

There was a founding event below/downstream of L151 (aka P310) some time around 3000 BC (broad margin of error).  P312 and U106 (and S1194) were born a few generations later and firm evidence of their existence is found in ancient eastern and central Bell Beaker burials in Europe dating from the early-to-mid-third millennium BC.  ZZ11 is a 'grandson' of P312.  The oldest R1b-P312+ burial so far might be I11443/BU32, Italy_Sicily_EBA, at 2872-2476 calBCE (c. 2674 BC) but the  original paper by Fernandes et al., 2020, gave a younger date (C14 dated to 2279-2102 calBCE).  The two contemporary burials, EHU002 (UE 450) 2562–2306 cal BCE (c. 2434 BC) El Hundido, Monasterio de Rodilla, Burgos, Castilla y León, Spain and I1389, dated to 4382 years before present (c. 2400 BC) from Sierentz, Les Villas d'Aurele, Haut-Rhin, France (Upper Rhine just north of the Alps) could therefore be the oldest examples of R1b-P312 yet found.  The oldest R1b-P312>ZZ11 so far, RISE563 (U152) from c. 2542 BC (2572-2512 calBCE), was also found just north of the Alps in Bavaria, Germany.

P312 Bell Beaker tribes, possibly with origins in Single Grave Culture in northwestern Europe, suddenly branched off in a large demographic expansion that had reached across the whole continent of Europe by c. 2500 BC.  The early DF27 and U152 Corded Ware/Bell Beaker clans were wide-ranging groups of paternally related males - descended from two founding events by two closely related men.  P312>Z40481>ZZ11>DF27>ZZ12 appears to be responsible for a big founding event.

The earliest burial with yDNA belonging to haplogroup DF27 is currently GBVPK (DF27>Z195) from the January 2021 paper Heterogeneous Hunter-Gatherer and Steppe-Related Ancestries in Late Neolithic and Bell Beaker Genomes from Present-Day France, Andaine Seguin-Orlando et al..  GBVPK is dated to 2461-2299 cal BC (c. 2380 BC).  The burial has high steppe-derived ancestry ('closer affinity Yamanaya steppe herder from Samara'): link.

Patterns of LD [Linkage Disequilibrium] decay provided an estimate for the time when steppe-derived genetic ancestry entered southern France at around ∼9.2 ± 3.6 to 10.4 ± 3.5 generations prior to the age of the earliest individual showing Yamnaya ancestry.

They estimate that the ancestor of GBVPK was in France in Corded Ware times c. 2650 BC - earlier than similar steppe-derived ancestry is found in Britain or Iberia.  Significant Corded Ware steppe influence was to the north and east of Europe at that time.  Z195+ GBVPK  with high steppe auDNA was found among four Bell Beaker and two Early Bronze Age individuals with no steppe auDNA from the Grotte des Tortues and Grotte Basse de la Vigne Perdue, both located in what is today the La Clape massif, southern France.

One of the next oldest DF27 burials, although there is not much difference considering the wide margins of error, is I0806 (.bam file analysed and identified as DF27+ by R. Rocca, 2016).  That was the first prehistoric DF27 burial to be identified and was a man with eastern/Baltic autosomal DNA who, according to Bell Beaker Blogger, was buried with a Corded Ware axe and a Bell Beaker c. 2290 BC (2431-2150 calBC) near the Bode river in what is now Quedlinburg, Germany.  I0806 is therefore potentially a representative of a people that M. Furholt has dubbed the 'SGBR' (Late Neolithic and Early Bronze Age Single Grave Burial Ritual Complex), another term for steppe-origin Bell Beaker people.  

SGBR appears in Central Europe and southern Scandinavia around 2900 BC, arrives on the British Isles a few hundred years later, and prevails until cremation burials take over, sometime after 1400 BC. Martin Furholt, 2019: link   

I0806's stone shaft-hole axe, displayed on Bell Beaker Blogger's website, has similarities with artifacts from the Catacombe Culture (c. 2800–2200 BC).  I0806 is one of the Bell Beaker/Corded Ware samples used by Haak et al. 2015 as evidence that Indo-European languages came from the steppes with R1b-M269>P312 men.  In 2016 the possibility that P312 subclades might be descended from a Corded Ware population was dismissed as unlikely.  Ancient DNA findings in 2019 suggested otherwise.  As with GBVPK in France, I0806's steppe autosomal ancestry was the highest of the Bell Beaker burials analysed in that area.

Identification of DF27 prehistoric burials had been missing from earlier ancient DNA studies due to lack of coverage in the tests used.  The technology used in SNP tests has difficulty identifying DF27 - even in modern samples.  Some of the undefined R1b-L151 (P310*) burials from studies could be DF27.  'NGS' (Next Generation Sequencing) testing does at least highlight the downstream SNPs and understanding is improving with the sharing of full NGS/WGS results on phylogenetic trees and analysis of ancient archaeological yDNA (archaeogenetics).  Unfortunately, tests used by archaeological studies (1240K panel) may not have full NGS/WGS coverage of the part of the y chromosome that DF27 and U152 are located at and often return only P312* or P310* results for potentially old and degraded DF27>ZZ12 archaeological remains.  The ISOGG 2019–2020 tree used by academia does not include many of the dozens of subclades downstream of DF27>ZZ12 that can be seen on FTDNA's Haplotree.

Lemercier map of Bell Beaker Civilization 2500-2200/2100 BCE, 2015-2017, above, from Bell Beaker Blogger

BBC TV documentary series, Invasion! with Dr. Sam Willis, broadcast 5th December 2017, covered Beaker DNA (around 20 minutes in).  They estimated that 70% of Britain's modern DNA traces back to the Beaker migration event 4500 years ago.  The presenter Dr. Willis states:

The Beaker migration originated from the steppes, from southern Ukraine and southern Russia.

Professor Ian Barnes of the Natural History Museum in London talks of the Beaker migration as being,

 perhaps the single-most significant migration event into the British Isles - the Beaker people seem to replace the Neolithic farmers.

They go on to say the Beaker invasion was the last major migration event that can be picked up using ancient DNA and there have been no major changes to Britain's overall DNA ever since.  The much later Iron Age/Celtic culture diffused language, innovations and fashions between those Beaker descendants in Europe and Britain.  Subsequent invasions of Britain and Ireland by Anglo-Saxons, Vikings and Normans came from peoples who were of the same Beaker origins as those already in Britain.  Towards the end of the episode Dr. Willis says:

the blood, or the DNA, of the Vikings and Anglo-Saxons was the same - they were descendants of the Beakers from the steppes.

A Yamnaya-like male dated to 2850–2500 BC has been found in Saxony-Anhalt, an area that has produced other early R1b Bell Beaker samples, link.  

Burials from the July 2019 paper, Ancient Genomes Reveal Yamnaya-Related Ancestry and a Potential Source of Indo-European Speakers in Iron Age Tianshan, Chao Ning et al., are R1b1a1a2 (R1b-M269).  R1b-M269 is a Bronze Age marker potentially linked with the expansion out of the steppes.  According to the summary, 'Our findings furthermore support a Yamnaya-related origin for the now extinct Tocharian languages in the Tarim Basin, in southern Xinjiang.'  However, according to Open Genomes, writing on Eurogenes, the two R1b burials are M15-2: R-PH200 and MO12: R-PH155.  These are ancient R1b branches that came about before R1b-M269 and are parallel lineages.

An important thing to note is that R-PH155 was found in a supposed "Gepid" from Serbia with an artificially deformed skull who was at least 20% East Asian, and therefore likely a Hun, and a Tian Shan Hun from Uzbekistan.

An important and fascinating early Iberian archaeological site is, PP4-Montelirio Structure 10.049, dated to 2850-2750 B.C at Valencina de la Concepción, Seville, southern Spain.  The site covers a huge area of land.  There is an elaborate burial containing exquisite crystal weapons (see below) and a mix of eastern, Moroccan and Iberian elements.  The large burial chamber housed a steppic-like steale and a number of gold, amber, African ivory and ostrich-derived objects.  The man was buried with around 24 women aged between 20 and 30.  Incidentally, a man with something like this number of wives might produce a large yDNA founding event (many parallel sons) like the one seen downstream of DF27>ZZ12.  The yDNA is thought to be being analysed and will hopefully be published: link.  There is no indication as to what his yDNA haplogroup is yet.

EARLY DF27

Bell Beaker man GBVPK (DF27>Z195, c. 2380 BC) in France was born several centuries after the formation of P312>Z40481>ZZ11>DF27 itself (DF27 was 'born' c. 3000 BC (4500 years ago + or - a margin of error, according to YFull).  DF27>Z195+ GBVPK had high steppe-derived autosomal ancestry.  DF27+ Bell Beaker/Corded Ware I0806 in Quedlinburg, Germany, c. 2290 BC, also had steppe ancestry and hailed from the northeast/Baltic (Eastern Hunter Gatherer/Baltic), not the west.  The well-travelled R1b-P310/L151* Boscombe Bowman and Amesbury Archer of Amesbury Down in southern England, of a similar date to I0806, are Bell Beaker burials with more southern/western Neolithic autosomal ancestry (also known as EEF or 'Early European Farmer') than other Bell Beaker men in Britain at that time.  Autosomal DNA comes in varying amounts from both the mother and the father of the individual.  Steppe ancestry men having children with European Neolithic women would produce offspring with less steppe admixture over the following generations.  It appears that as DF27 males migrated further south and west and they married into existing local populations their EEF autosomal ancestry increased and their steppe ancestry decreased over time.  Individual males that then migrated north, like the Bowman and Archer, subsequently stand out in archaeogenetic analysis when found buried among a population that contained little Neolithic EEF ancestry.

Eastern R1b-P312 Bell Beaker men with steppe ancestry probably arrived at their most westerly destinations, Britain and Iberia, in many ways.  Bell Beaker/Corded Ware people were traversing all of Europe, trading and bringing innovations like metal working to some areas.  As mentioned, those eastern Bell Beaker males belonged to different y haplogroups (P312) to the existing Neolithic farmer groups.  According to Harvard geneticist David Reich, the displacement of local males in the British Isles by P312 Bell Beaker men after c. 2500 BC was very swift.  Others think the process might have taken longer.  In Iberia the process may have taken about 500 years longer.  Iberia could have had a much bigger surviving population than the British Isles did at the time of P312's arrival.  Metal working was already known in Iberia - it might not have been as much of an advantage for the newcomers as it was in Britain.  These changes occurred between c. 2500-2200BC when new elements from the east arrive in the second half of the Bell Beaker era in Iberia.  It is suggested here (Kristiansen 2018 Vienna Genes talk, 2018) that a Neolithic plague, one of the first incidences of what would be known later in medieval times as the Black Death, reduced the population of Europe prior to the arrival of hostile steppe males from the east in 3000 BC, link.   However, the preexisting female population of Iberia appears to have largely survived the arrival of R1b-P312, unlike in Britain where many of the previous Neolithic inhabitants disappear.  Ancient genome-wide DNA from France highlights the complexity of interactions between Mesolithic hunter-gatherers and Neolithic farmers, Maïté Rivollat et al., May 2020, link, highlights the population turnover.  In 2200 BC there was a prolonged period of drought (4.2-kiloyear event) that lasted for a century and caused worldwide population collapse, link.  On the Iberian Peninsula this climate change led to the construction of motillas-type settlements to access ground water.

As mentioned, U152 and DF27 share the same 'father', ZZ11.  U152 RISE563, from what is now Osterhofen-Altenmarkt in Germany, belongs to 'Bell Beaker East' and dates to c. 2542 BC (2572-2512 calBCE).  He was a migrant, based on isotope analysis, and autosomal DNA plots with Corded Ware and modern day eastern Ukrainians, Kargopol Russians and Mordovians.  DF27>ZZ12 is today geographically widespread.  The latest ancient yDNA papers all conclude that the P312 remains they tested may have had origins further to the east of Europe.  The abstract of the paper by Iñigo Olalde and David Reich, 2017 concludes:

The arrival of the BBC in Britain can thus be viewed as the western continuation of the massive movement of people that brought the Corded Ware Complex and steppe ancestry into central Europe a few hundred years before.

Members of the Single Grave Culture buried their dead with stone battle-axes.  R-L51, the ancestor of P312>ZZ11, probably originated somewhere in Corded Ware territory.

There was discussion at Anthrogenica in January 2019 suggesting a potential link between descendants of the Corded Ware culture in the west (Single Grave Culture c. 2800 BC) and a founding event of R1b-P312 and expansion across all northwestern Europe of its downstream subclades from Dutch Bell Beaker in the region of the Lower Rhine.  All the samples from the Oostwoud-Tuithoorn site in what is now West Frisia are said to have similarly high steppe autosomal ancestry.  R1b-P312* Beaker man, I5748 , dated to c. 2400 BC (2579–2233 calBCE) is one of the oldest R1b-P312 burials in Europe.  He had a 'no call' for U152 and DF27 downstream of P312.  In March 2021 Alex Williamson identified I5748 as being DF19>Z302.  Two of the Oostwoud-Tuithoorn Dutch Bell Beaker samples, the only ones that had data at DF27's position, I4074 dated c. 2096 BC (2278–1914 calBCE) and I4073 from c. 2015 BC (2196–1903 calBCE), were P312* and negative for DF27.  Apparently, all Bell Beaker period dates are uncertain and have wide margins of error ('fluctuations in atmospheric C14 fractions ').  Another interesting transitional Corded Ware/Bell Beaker grave in The Netherlands is mentioned here.

IBERIAN BELL BEAKER

Early results from Parallel ancient genomic transects reveal complex population history of early European farmers, March 6, 2017 showed the earliest Iberian 'Bell Beaker' period people were not R1b but haplgroup I and I2a2a and had no eastern/steppe autosomal DNA.  This apparent I2 dominance in pre-Beaker Britain and Iberia has been confirmed by more recent findings in the large March 2019 Olalde Iberian study.  In 9000 BC I2 farmers spread from the Fertile Crescent and Anatolia across the Mediterranean, reaching Britain via Atlantic Iberia and France by around 4000 BC.  See Ancient genomes indicate population replacement in Early Neolithic Britain, in Nature, April 2019 and also BBC Science article: link.  The three 'El Sotillo' remains date to between 2916 BC - 2212 BC and were found in a collective megalithic tomb that contained remains of Bell Beaker pottery.  The oldest 'Bell Beaker' period skeleton to be yDNA tested anywhere so far, ES-6G-110, was found there.  He was I2a2a and dates to 2916-2714 BC.  Those Neolithic Iberian people in collective megalithic tombs are different to later eastern Bell Beaker people (P312 with steppe admixture) who moved into Iberia and were buried individually or 'individualised' under burial mounds.  Nature article Y-chromosome target enrichment reveals rapid expansion of haplogroup R1b-DF27 in Iberia during the Bronze Age transition, Carla García-Fernández et al., December 2022, focuses on DF27 in Iberia but uses the ISOGG 2019–2020 tree.  The many DF27>ZZ12 subclades discovered by Alex Williamson's Big Tree in 2014 and later represented on the FTDNA Haplotree (the DF27* paragroup in the paper) are not mentioned.  The oldest-known ancient DF27 burials (see below) are not listed.  The authors tentatively suggest an origin in northeast Iberia for DF27:

The estimated age was highest in the Basque Country, Catalonia and NW Europe, and lowest in Central Spain. However, differences are small and cannot be used to pinpoint a place of origin for R1b-DF27, since it is likely that the initial Bronze Age event carried R1b-DF27 throughout Iberia within a short timeframe.

The present results show indeed that nucleotide diversity is marginally higher in NE Iberia than elsewhere, and that whole branches of the R1b-DF27 phylogeny, particularly R1b-Z272, are restricted to Iberia. However, ancient DNA, and the fact that nucleotide diversity is not significantly lower in NW Europe compared to Iberia, do not rule out the possibility that R1b-DF27 originated elsewhere in Western Europe, but expanded and radiated in the north of the Iberian Peninsula, where it replaced the local paternal lineages to a great extent.

A Spanish study, due for release, (El Argar culture and the ancient sites of La Bastida, c. 2000 BC and La Almoloya, 2200-1550 BC) found that some 'outsiders' may have arrived there from south of present-day Russia.  PIR001 El Pirulejo (Priego de Córdoba) is R1b-M269 (pdf):

The burial was accompanied by a low angle vase which typologically was related to similar forms of the Argar culture (2100-1450 BC) of the Iberian southeast.

In 2017 a native Spanish speaker translated a video presentation thus: 

They advance that the few males they have from the sites of La Bastida and La Almoloya, they show that males came from outside, originating in the P-C steppe, moving to Western Europe, British Islands and from there arriving to Galicia (NW Iberia) and finally reaching the South East, where El Argar culture is placed.

C14 dates the wall of La Bastida to 2200 BC.  The design of the wall has no parallels in the rest of Iberia or Western and Central Europe.  The closest resemblance to the design of La Bastida is in Troy and other places in the Levant.  

In this system, La Bastida was a central point which needed special protection. If La Bastida was the capital of the whole El Argar territory or one of a series of regional centers is difficult to say at the present moment. Whatever the case, it was the largest eco-nomic and political center to be established immediately after the 2200 cal BC abandonment or destruction of most Copper Age settlements., link 

DF27>Z195+ ALM039 is the male burial from a high status Argaric Bronze Age (3rd phase) grave (Grave 38) at La Almoloya, Murcia, Spain (1740 - 1533 cal BC, lab code MAMS-22231).  He was buried under the floor of a room in a large ovoid jar with a woman wearing the silver diadem, below, in possibly the earliest Bronze Age palace yet discovered in western Europe.  The man and woman died at around the same time in the mid seventeenth century BC and their estimated ages at death were 25–30 and 35–40 years respectively.  The man was buried with gold items, the woman with fine silver ones.  The room that holds their tomb, named H9, looks like a hall designed for social gatherings with benches around the plastered stone walls and a raised podium near a large hearth.  Link.

Above: Silver diadem from La Almoloya grave 38 (figure by J.A. Soldevilla, courtesy of the Arqueoecologia Social Mediterrània Research Group, Universitat Autònoma de Barcelona).

Genomic transformation and social organization during the Copper Age–Bronze Age transition in southern Iberia, Vanessa Villalba-Mouco et al., 2021, link, states in the Abstract:

Concomitant with the rise of El Argar starting ~2200 cal BCE, we observe a complete turnover of Y-chromosome lineages along with the arrival of Steppe-related ancestry. This pattern is consistent with a founder effect in male lineages, supported by our finding that males shared more relatives at sites than females., link

The beginning of the Bronze Age [Early Bronze Age (EBA)] in Iberia (2200 to 1550 cal BCE) marks a clear population turnover, suggested by both the omnipresence of steppe-related ancestry in all individuals directly postdating 2200 BCE. An even more notable shift can be observed in the frequency of Y-chromosome haplogroups in males, which are almost exclusively of the R1b-P312 type that was completely absent in Iberia before 2400 BCE.

There appear to be many DF27 burials with steppe-related ancestry identified in that paper that beong to the El Argar culture of c. 2200 BC to c. 1500 BC.  The previous Copper Age inhabitants of Iberia had no steppe-related ancestry.  From the paper:

The resulting f4-values confirmed a smaller amount of steppe-related ancestry in individuals from the Argaric sites La Almoloya and La Bastida compared to the rest of Iberia_BA groups, especially when compared to those from northern Iberia (fig. S5B and table S2.8), despite the complete turnover to lineage R1b-P312 (except for one subadult male in La Bastida) visible in the Y-chromosome record (Fig. 3B, table S2.6, and text S8).

Most burials that could be identified to a higher resolution are Z195 but many remain P312* and therefore some may be as-yet-unrecognized DF27>ZZ12 subclades.  R. Rocca found in a preliminary scan that here are 17 Z195+ burials, 7 P312* burials and four others that are P312>Z40481>ZZ11, L51, P311 and L151 from La Almoloya, the large El Argar settlement located in the northern foothills of Sierra Espuña mentioned above.  There are 6 as-yet-undefined P312* burials at La Bastida:

a hilltop settlement that was naturally protected by vertical slopes and a monumental fortification in a mountainous topography of the core area of El Argar formation.

None of the La Bastida burials were found to be DF27>Z195.  The oldest P312 burial at La Bastida was:

BAS023 (Arc. ID: BA88): Adult male burial directly dated to 2122-2093 cal (c. 2107 BC).

The earliest R1b-P312 burial in Iberia is EHU002 (UE 450) 2562–2306 cal BCE (3933±32 BP, CSIC-1896) El Hundido, Monasterio de Rodilla, Burgos, Castilla y León, Spain.  At c. 2434 BC, EHU002 is younger than P312>U152 RISE563, dated to 2572-2512 calBCE (c. 2542 BC), although considering the wide margins of error there is not much difference.  Both are probably a few hundred years younger than the first U152 and DF27 'brothers'.  EHU002 is P312+ DF19- L238- Z290- L21- but it is not known if he is + or - for DF99, DF27, Z195, ZZ12, U152 or L2.  As with other R1b-P312* and R1b-P310* results (the asterisk meaning no downstream SNPs found) from time-degraded ancient DNA, and considering the location of the burial not far from oldest DF27+ burial GBVPK at Grotte Basse de la Vigne Perdue ( c. 2380 BC), there is a good likelihood that EHU002 is DF27+ but unfortunately there is no SNP evidence.

Corded Ware and Bell Beaker cultures covered a large area and isotopic analysis of the teeth of Bell Beaker burials has revealed that some individuals were well-travelled.  The R1b-P310* Amesbury Archer and Boscombe Boman were foreigners with 'southern' autosomal admixture buried in prehistoric England, near Stonehenge (c. 2300 BC), for example.  Eastern Bell Beaker (those originally from the east with steppe/Corded Ware ancestry) may have arrived in Iberia between 2500 and 2000 BC, bringing R1b-P312.

Regarding DF27 in The genomic history of the Iberian Peninsula over the past 8000 years, Iñigo Olalde et al., March 2019, Supplement:  

From the Bronze Age (~2200–900 BCE), we increase the available dataset (6,7,17) from 7 to 60 individuals and show how ancestry from the Pontic-Caspian steppe (Steppe ancestry) appeared throughout Iberia in this period (Fig. 1, C and D), albeit with less impact in the south (table S13).  The earliest evidence is in 14 individuals dated to ~2500–2000 BCE who coexisted with local people without Steppe ancestry (Fig. 2B ).  These groups lived in close proximity and admixed to form the Bronze Age population after 2000 BCE with ~40% ancestry from incoming groups (Fig. 2B and fig. S6). Y-chromosome turnover was even more pronounced (Fig. 2B ), as the lineages common in Copper Age Iberia (I2, G2, and H) were almost completely replaced by one lineage, R1b-M269.

The Arrival of Steppe and Iranian Related Ancestry in the Islands of the Western Mediterranean, Daniel M Fernandes et al., March 2019 states:

The presence of Steppe ancestry in Early Bronze Age Sicily is also evident in Y chromosome analysis, which reveals that 4 of the 5 Early Bronze Age males had Steppe-associated Y-haplogroup R1b1a1a2a1a2. (Online Table 1).  Two of these were Y- haplogroup R1b1a1a2a1a2a1 (Z195).

The other individual, the one without 'Steppe-associated' yDNA in the EBA Sicily group, was haplogroup J.  The two DF27>Z195 males date to around 2200 BC and are therefor not quite as old as GBVPK in France (c. 2380 BC) or I0806 in Germany (c. 2290 BC).  The two DF27>Z195 males are I8561, from Isnello (Abisso del Vento), (2346-2199 calBCE) and I3123 from Buffa Cave near Villafrati, (2287-2044 calBCE).  I3123 is said to be extremely brachycephalic (planoccipital) meaning a round, broad skull flattened at the back - a characteristic Bell Beaker head shape seen in other northern and eastern European Bell Beaker burials.  The two R1b-P312, burials, both from Buffa Cave, are I11443 (Petrous bone) with highest 'steppe ancestry' (40.2 ± 3.5%) C14 dated to 2279-2102 calBCE and I11442 (Temporal bones) C14 dated to 2281-2047 calBCE.  Both were only found to be P312xR1b1a1a2a1a2c (P312xL21), so they were not tested for DF27 (ZZ12 etc.) itself.  The Z195 individuals also have high steppe ancestry: I8561 (23.3 ± 3.5%), and I3123 (14.1 ± 3.4%), link.  I11442 is P312+ and from Buffa Cave, dated to 2281-2047 calBCE with possible Iranian-related (from Anatolia) ancestry.  This might be due to low coverage of the ancient DNA or it might be real - there has been Iran-Caucasus ancestry found in Neolithic Italy.  

Early Bronze Age Sicily: Among the Sicilian Early Bronze assemblage individuals, I11443 from Buffa Cave (Sicily_EBA11443) and individual I8561 from Isnello (Sicily_EBA8561) were not consistent with forming a clade with the remaining Early Bronze Age Sicilians in the great majority of tests.  We therefore treat them as separate outliers (Supplementary Table 5).

R1b-P312>? I11443's ancestry can be modeled as Sicily Middle Neolithic in origin mixed with 74% French Bell Beaker. Likewise, DF27>Z195 I8651 had 45% French Bell Beaker and I3123, the DF27>Z195 individual with the Bell Beaker head shape, was Sicilian Middle Neolithic with 30% French Bell Beaker, link.  If DF27 reached Sicily so early it is likely that Bell Beaker people could have known the entire Mediterranean as early as 2200 BC, although there is not much evidence in the archaeological record.  The Villalba-Mouco et al., 2021, paper found that El Argar DF27 burials have steppe-related ancestry but have slightly less of it than those in the north of Iberia and northern Europe.  They suggest Mediterranean and central European ancestries shaped the genetic profile of southeastern BA groups like El Argar in Iberia.  Some of those southeastern Iberian Bell Beakers have Anatolian and Aegean-related admixture and have some ancestry from people belonging to ancient Mediterranean cultures like the Minoans and Mycenaeans.  The Villalba-Mouco et al. paper also found that burial ZAP002 (Z195>Z296>Z268>A7066>BY32727):

is highly likely a non-local individual, who carries steppe-related, central Mediterranean and North African ancestries.

There is an Italian Bell Beaker individual much further north, I2478 from Parma (2200–1930 calBCE).  He is also R1b-P312* like two of the Sicilian Bell Beakers and he owned a 'remarkable' lithic knife.  He is negative for U152 and might potentially be DF27.  

Marie Besse's map of Bell Beaker collective graves versus Bell Beaker single graves, above.

NORTHERN BELL BEAKER

I0806 in Germany was buried in around 2290 BC on his own (single grave) with a Corded Ware battle axe and had eastern/steppe autosomal DNA.  This region was occupied by the Corded Ware culture (c. 2900 - c. 2350 BC).  Corded Ware covered a vast territory and was contemporary with Bell Beaker.  The two cultures overlapped in Germany, where I0806 was buried.  The Corded Ware area also coincides with the distribution of 'eastern' Bell Beaker arrowheads.  Corded Ware culture is known as Battle Axe culture in Scandinavia.  An early civilization that ruled Orkney and built the Ness of Brodgar temple complex (c. 3200 BC-2200 BC) may be key to the story of the exploration of the north by Bell Beaker people in the Neolithic and Bronze Age.  A study, Continental migrants and Neolithic male lineages in Bronze Age Orkney, University of Huddersfield, 2022, addresses some of the questions:

Unlike elsewhere in Britain, there is little material evidence to suggest a Beaker presence, suggesting that Orkney may have developed along an insular trajectory during the second millennium BC ..uniquely in northern and central Europe, most of the male lineages were inherited from the local Neolithic.', link

Ancient DNA at the edge of the world: Continental immigration and the persistence of Neolithic male lineages in Bronze Age Orkney, Katharina Dulias et al. was published in February 2022, link.  Significant Bronze Age steppe ancestry on the islands looks to have arrived with women.  Although they caution that the samples were from a peripheral part of the islands and more data is required from mainland Orkney, it appears that the males on Orkney 'held out' against the Bell Beakers for around 1000 years longer than the rest of the British Isles did in the Bronze Age.  The paper Imputed genomes and haplotype-based analyses of the Picts of early medieval Scotland reveal fine-scale relatedness between Iron Age, early medieval and the modern people of the UK., Adeline Morez et al., 2022, link, examines the ancestry of the Picts of early medieval Scotland (ca. AD 300-900).

The Iron Age and unadmixed Viking Age Orcadians also show the highest degree of a red ancestry component (Fig 2B, S7 Fig), which is inconsistent with having originated from direct gene flow from any population included in this study and instead likely reflects retention of a less diverse pre-Iron Age ancestry in Orkney and/or strong genetic drift (such as a bottleneck or founder effect). In fact, recent research show that Bronze Age populations in Orkney were differentiated from their counterparts on mainland Britain due to retention of male Neolithic ancestry (Y-chromosomal haplogroup I2), while the R1b haplogroup associated with Bell Beaker expansion largely replaced the I2 haplogroups in the rest of Britain, implying that local ancestry may have persisted also into the Iron Age and early medieval period.

Late Copper Age migrations can be seen on this map at indo-european.eu. It includes the oldest DF27 located so far, around Mittelelbe-Saale (pale blue symbols in Germany).  The arrival of Bell Beaker in Jutland, the Danish islands and southwest Norway c. 2350 BC triggered the onset of the Late Neolithic period in Scandinavia - a prelude to the Nordic Bronze Age.  Population Genomics of Stone Age Eurasia, Morten E. Allentoft et al., 2022, link, suggests that R1b-P310 lineages appear in southern Scandinavia until around 3800 BP.  That time, c. 2000 BC, coincides with the expansion of Bell Beaker.  They say that R1b-P310 were preceded by R1a in the Corded Ware period (4600 - 4300 BP) and followed by the arrival of I1 after 3800 BP.  

Iberia was an important source of precious commodities, particularly copper and gold.  Metal ores and metalworking skills were imported into Scandinavia in the Bronze Age from Britain and Europe.  The Danish study, Mapping human mobility during the third and second millennia BC in present-day Denmark, Karin Margarita Frei et al., August 2019, suggests that new people moved in to the area at the start of the Nordic Bronze Age:

 our data also indicate a clear shift in mobility patterns from around 1600 BC onwards, with a larger variation in the geographical origin of the migrants, and potentially including more distant regions.  This shift occurred during a transition period at the beginning of the Nordic Bronze Age at a time when society flourished, expanded and experienced an unprecedented economic growth, suggesting that these aspects were closely related., link

The shift in human mobility characterized by the expansion in diversity of areas of origin of the non-locals appears to occur parallel to the emergence of the long-distance metal trade that connected present-day Denmark to areas in, e.g., central and southern Europe as well as the British Isles.

In Population genomics of the Viking world, Ashot Margaryan et al., 2019 Nordic Bronze Age and Early Viking Age ancient DNA obtained from burials shows an increase in 'southern' autosomal DNA.  Traders and warriors from the south travelled along the Atlantic maritime trade routes in long boats, as depicted in Scandinavian rock art.  From The ‘Stranger King’ (bull) and rock art, Johan Ling and Michael Rowlands:

In fact, several Scandinavian features from the Bronze Age have been inspired by the Mediterranean (Thrane 1990; Kristiansen and Larsson 2005) and there are some striking similarities between the horned warriors in Extremaduran rock art, Nuragic figurines from Sardinia, depictions of ‘sea peoples’ with horned helmets in the tomb of Ramesses II and the horned warriors depicted on the rock art in western Sweden and the horned bronze figurines from Grevesvänge in Denmark (Fig. 8.10, see also Harrison 2004; Kristiansen and Larsson 2005). In this context it is important to stress that Scandinavia also ‘imported’ copper from these regions during the Bronze Age (Ling et al. 2012). Even if these horned anthropomorphs were produced in remote regions during the Bronze Age, they co-exist within, more or less, the same epoch, 1200–800 BC (Harrison 2004).

NORTH AND SOUTH?

Iberia, the Atlantic Bronze Age and the Mediterranean, Brendan O'Connor et al., 2008:

Precocious activity on the Atlantic sea routes in the Copper Age is suggested by diffusion of the Beaker phenomenon and its associated «package» of artefacts, notably its metal such as Palmela points, and also by the spread of early goldwork fashions such as lunulae.

Throughout the Early and Middle Bronze Age long distance movement along the Atlantic seaways appears patchy and slight. It was perhaps because of this isolation that Iberian metal working remained conservative throughout the Middle Bronze Age, the range and sophistication of its products slight compared with Atlantic regions further north.

last gasp of the Iberian Bronze Age saw some of the most remarkable long-range traffic linking the Atlantic and Mediterranean worlds., link

The Bronze Age in Iberia began c. 1800 BC, about 600 years after Bell Beaker men with steppe ancestry, R1b-DF27 GBVPK and R1b-P312* EHU002, were buried in southern France and northern Spain respectively.  DF27>ZZ12>Z46512>FGC78762>ZZ19 burial I15033 [listed as P312 in the Patterson et al. 2021 paper but Alex Williamson found ZZ19] at Port Blanc, Brittany dates to 1891–1743 cal BCE (c. 1817 BC).  The site of Port Blanc is a megalithic complex in Brittany, in the north-west of France.  ZZ19 is a large prehistoric subclade that dates back to Bell Beaker times.  This is currently the oldest ancient example of ZZ19 to be found.

Atlantic traders were taking copper ore from Iberia and tin from Cornwall in Britain as far as Sweden for use in the emerging Bronze Age cultures in Scandinavia.  Sea routes along the Norwegian coast were travelled in the Late Neolithic and Bronze Age and settlements and burial mounds appear there at that time.  From 2000 BC - 1400 BC Britain was also an important source of copper.  The scale of prehistoric mining at the Great Orme mine in North Wales was huge, they dug out enough malachite to make 11 million bronze axe heads.  The Welsh copper mine is the biggest prehistoric man-made underground void in the world.

Baltic amber from the north travelled south to the Mediterranean through the land and maritime trade networks.  From the Bronze Age onward there was radical material and social transformation in Iberia.  Before then, from at least the fourth millennium BC, Sicilian amber (simetite) was imported into Iberia from the south across the Mediterranean sea route.  After a decline in that trade new significant amounts of northern Baltic amber appear in Iberia from c. 2000 BC onward (Settlement patterns in the central and southeastern area of Iberian peninsula, link).  Evidence of these wealthy long distance travellers comes in the form of a teenage boy who was determined, through tooth isotope analysis, to have origins in the Mediterranean region.  He was buried with a Baltic amber necklace and his burial is dated to 1550 BC at Boscombe Down, near Stonehenge in Wiltshire, England.  More on the Boscombe Bell Beaker burials on the Bronze Age DF27 page.  A tanged bronze dagger, possibly made in Cyprus and thought to date to around 2000 BC, was found a long time ago by an early antiquarian (no proper context) on Egton Moor in northeast Yorkshire.

Map above showing possible flows and routes of metal from the mining districts in Europe to Scandinavia in the Bronze Age.  From The ‘Stranger King’ (bull) and rock art, Johan Ling and Michael Rowlands (pdf)

The place that the large modern R1b-P312 subclades are found in roughly equal numbers is central/northwest France.  Some early DF27 subclades could have entered both Britain and Iberia from earlier Neolithic origins in Corded Ware/Single Grave Culture territory in north/west/central Europe.  The 'star burst' expansion of DF27 Bell Beaker subclade founders looks to have happened in the Bronze Age and the movement of Atlantic Bronze Age maritime peoples might be a factor in the 'North-South' geographical split seen in descendants of the many parallel DF27 subclades on the phylogenetic trees.  Iberia, the Atlantic Bronze Age and the Mediterranean, Brendan O'Connor et al., 2008:

With the onset of LBA2/BF2 and the turn of the twelfth-eleventh centuries BC, everything changed in Atlantic Iberia. Suddenly there are abundant signs of Atlantic metalwork and metalworking influence, as well as numerous hoards, castros and Hohensiedlungen. Only now can one in any sense speak of an Atlantic Bronze Age, and for the first time see metal production with common echoes all the way from northern France and Britain to southern Spain. But connections were not only with the north and the Atlantic world. On the sites and in the hoards, Atlantic elements are mixed with eastern material coming from the central Mediterranean and beyond (Burgess 1991)., link

1500-1100 BC was the period of extensive interaction between people along the Atlantic coast up into Scandinavia.  Seafaring warriors with new Mediterranean ship technology took large and valuable cargoes of metal ore along the Bronze Age maritories from Iberia up along the northward maritime trade route while Baltic amber was taken south.  Ores could be exported to the north in greater quantities by ship - the transport of the first Scandinavian-bound copper ore from its source north of the Alps involved difficult overland travel with smaller cargoes.  Perhaps difficulties with the transportation of precious goods overland led to a switch to maritime trade routes at that time.  The finer analysis of specific subclades below DF27 is required to determine who was moving where in prehistory.

The context, phylogeny and dating of ancient archaeological yDNA remains with the identification of specific subclades in Iberia and Britain will be crucial.  In a few years there could be tens of thousands of ancient genome sequences and the reference material from the phylogenetic trees will be more accurate.  The autosomal DNA of all northwestern Europeans is very similar and probably has been so since the Bell Beaker period.  There is little difference in autosomal DNA results between the 'Celtic' people in the west of the British Isles and Ireland and 'Germanic' people around the North Sea even though there is variation in the frequencies of yDNA haplogroups in those different regions.  A PCA (Principle Component Analysis), made by Eurogenes blog comparing modern northern Europeans with Bell Beaker burials in Olalde et al. 2018, shows Bell Beaker plots (has closest relatedness) with both Germanic and Celtic present-day populations.  The PoBI (People of the British Isles) project found more genetic variation in the traditionally 'Celtic' west of the Isles than the more 'Germanic' east - indicating that the story was already complex before the Migration Period.  Studies show that most people in the British Isles, whether they identify as Celtic or Germanic, descend from the same Bell Beaker ancestors who were in Europe around 4500 years ago.  Bronze Age Britons had the same general Bell Beaker ancestry as later Anglo-Saxon, Viking and Norman immigrants.

Above, model of a Nuragic ship, Sardinia, by Shardan, link

Modern nation states and their accumulated historical baggage did not exist 5000 years ago.  Europe was a comparatively sparsely-populated territory through which people moved around on track ways, rivers and sea routes.  We might never know what language people belonging to the various individual ancient yDNA haplogroups spoke initially - prehistoric languages from the time of the Bell Beaker Culture are unattested.  See Reconciling material cultures in archaeology with genetic data: The nomenclature of clusters emerging from archaeogenomic analysis in Nature, link. 

Phylogenetic trees made up of present-day NGS tests show that the connection (a yDNA subclade's node or branch point) between most parallel lines in the phylogeny of DF27 converge in the Bronze Age.  The earliest Bell Beaker in Britain arrives in Wessex c. 2375 BC and in Scotland c. 2350 BC.  The Atlantic façade along which Maritime Beaker people travelled was huge, much bigger than the Atlantic coast itself, and included Sweden, Denmark, North Germany, The Netherlands, Belgium, Scotland, England, Wales, Ireland, northwest France, northern Spain, Portugal and the Mediterranean - parallel DF27 subclades voyaged between the Mediterranean and Nordic regions via the Atlantic Bronze Age maritime networks.

Metal-poor Scandinavia’s sustained demand for resources led to a prolonged symbiosis with the Atlantic façade and Central Europe during the Bronze Age. Complementary advantages of the Pre-Germanic North included Baltic amber and societies favourably situated and organized to build seagoing vessels and recruit crews for long-distance maritime expeditions. An integral dimension of this long-term network was intense contact between the Indo-European dialects that became Celtic and those that became Germanic. The Celto-Germanic vocabulary—like the motifs shared by Iberian stelae and Scandinavian rock art—illuminates this interaction, opening a window onto the European Bronze Age.  CELTO-GERMANIC Later Prehistory and Post-Proto-Indo-European vocabulary in the North and West, John Koch, University of Wales, 2020, Abstract,  link

Studies of isotopes in ancient remains, like that those of the early Bell Beaker migrants the Boscombe Bowmen and the Amesbury Archer buried near Stonehenge, Wessex in 2380–2290 cal BC, as well as Bee Low man, pictured below, buried in Derbyshire in a round barrow with a bronze pin and two possible awls but no pot in 2200–2030 cal BC, have shown that some early Bell Beaker individuals travelled far-and-wide across Europe in their own lifetimes.  Within Britain itself Bell Beaker people moved widely around the countryside.  See Beaker people in Britain: migration, mobility and diet, Mike Parker Pearson et al. 2016.  Given this high mobility within Bell Beaker society, it is possible that a founder from an early 4500 year old yDNA haplogroup could theoretically have settled and become established in any area of Bronze Age Europe.

LANGUAGES

Notions of Celtic and Germanic in Britain are relatively recent constructs - the people of northwest Europe descend largely from Bell Beaker ancestors living in the Late Neolithic period thousands of years before Celtic and Germanic linguistic groups are first attested.  The terms Germanic and Celtic used here refer to prehistoric people speaking early dialects.  Most present-day European languages developed from dialects of early Indo-European.  It is thought that people bearing R1b-P312 subclades brought Indo-European languages to Europe.  There are arguments that Bell Beaker people might or might not have spoken an Indo-European language but there is no evidence for any particular languages being spoken.  Early far-flung interrelated Bell Beaker groups cooperated across vast distances and it seems likely they originally spoke the same 'lingua franca' with perhaps some regional dialectical variations developing over time.  This early language shared vocabulary that would later be incorporated into Germanic and Celtic.  The 'Nordwestblock' theory proposes that a prehistoric culture in northwest Europe (present-day Netherlands, Belgium, northern France, and northwestern Germany) spoke a language that was neither Germanic nor Celtic but another language that existed somewhere in between.  The middle Bronze Age Hilversum culture, located in the region of modern Belgium, southern Netherlands and northern France, a culture that was related to Wessex culture in England, may have spoken this intermediate Nordwestblock language.  There is, and always has been, much scholarly debate about the origins of languages.  Indo-European language tree: link  

Proto-Germanic language is attested just a few hundred years before the birth of Christ when what became the Jastorf culture in the northwest of Europe lived next to La Tène Celts.  Greek: Keltoi and Latin: Celtae was a term that Graeco-Roman ethnographers applied to tribes living in the region of the upper Rhine and Danube during the sixth-to-first centuries BC.  Celtae became a term for people living west of the Rhine.  Germani was a term used by the Romans to describe all tribes living in the vicinity of the Rhine Delta (what is now Belgium, the Netherlands and Germany) around 2000 years ago.  After the Roman period the term fell into disuse.  Harry Fokkens, 2001, considered the area a 'border zone' with elements of the Atlantic maritime world mixing with the Nordic world.  Tribes labelled as Germanic or Celtic by outsiders might not have recognized or described themselves by that term - it had no emic significance to them.  The modern perception of a dichotomy between Celtic and Germanic may not have been pronounced around the North Sea basin in the Late Bronze Age/Iron Age, some vocabulary was shared.

Areas that eventually gave rise to the Germanic and Celtic languages may have contained a complex patchwork of neighbouring peoples speaking a variety of early dialects, all interacting, trading with and influencing each other to some extent.  The elites/chieftains that controlled the many nodes along the maritime networks probably belonged to a variety of differing yDNA lineages by the Iron Age but their autosomal DNA remained similar to Dutch Bell Beaker people buried thousands of years earlier.  When the earliest Runic writing to survive was carved c. 150 AD Bell Beaker man I0806 lying in a burial mound in Germany, and GBVPK in a cave in France, were ancient history.  The descendants of the Bell Beaker people were moving extensively around Europe for over 2000 years before modern language groups are first attested.  Languages spoken by representatives of more recent yDNA subclades might be inferred through their location, autosomal admixture and the historical and cultural context of their burials and grave goods, but it is unlikely that the obscure origins of unattested prehistoric linguistic groups will be defined by genetics.

Above, ship rock art, Tanum, Sweden, Public Domain (right), link

In around 2500 BC early R1b-P312 Bell Beaker men may have spoken an early form of Indo-European, i.e. 'North-West Indo-European'.  That early language is ancestral to Celtic, Italic, Germanic and Balto-Slavic languages.  More northerly Beaker groups once shared Celtic-Germanic vocabulary relating to war, ritual and horsemanship - words that were not shared by their early Beaker Italic-speaking relations.  The earliest embryonic forms of Celtic and Germanic might have origins in dialects of the late Bronze Age, at around the same time that imported metals and the horned helmet and ship iconography on rock art appeared in the North.  Celtic language is recorded as existing in the mid-to-late-first millennium BC, slightly earlier than Germanic is first attested.  The Celtic lexicon also shows early contact with Proto-Germanic and it has been suggested that Germanic was influenced by Celtic, as well as Roman and an ancient preexisting non-Indo European language.  Proto-Germanic contains loans from Celtic that followed Grimm's and Werner's laws, indicating that contacts took place before Proto-Germanic became linguistically defined.

From In Search of the Indo-Europeans by James Mallory, 1991:

it is most probable that the Corded Ware horizon provided the vector for a series of Indo-European languages that spread both to the west as far as Holland and east into the Baltic and upper Volga.  Out of this later emerged possibly the Celtic and Italic, and more certainly the Germanic, Baltic and Slavic languages.

Bronze Age and Iron Age groups were mobile. 

Human history is full of dead ends, and we should not expect the people who lived in any one place in the past to be the direct ancestors of those who live there today.  Who We Are and How We Got Here, David Reich, 2018.

EARLIEST DF27 (ISOGG's R1b1a1b1a1a2a) SO FAR  IDENTIFIED BY ARCHAEOGENETICS - Oldest to Youngest (bear in mind wide margins of error :)

Accurate identification of DF27 requires high resolution NGS/WGS testing and up-to-date SNP reference tables.  Many ancient yDNA samples have deteriorated with age and have missing or conflicting SNP information, so no ancient result can be regarded as being 'set in stone'.  Most early studies used lower-cost SNP arrays that only test for a small number of P312 SNPs and do not detect DF27.  Note the caveats.  

As mentioned above, EHU002 (UE 450) 2562–2306 cal BCE (c. 2434 BC) El Hundido, Monasterio de Rodilla, Burgos, Castilla y León, Spain with 'steppe ancestry' is P312+ DF19- L238- Z290- L21- but DF99, DF27, Z195, ZZ12, U152 and L2 are missing/no-calls.  As with other R1b-P312* and R1b-P310* results from time-degraded ancient DNA there is a good likelihood of EHU002 being derived/positive for DF27 but the SNPs are not present in the sample.  The high 'EEF' or 'southern' autosomal admixture R1b-P310* Boscombe Bowman from c. 2330 BC in Wessex, listed below, has an additional hint that he might be DF27+ in the form of FGC11381.  That SNP is potentially old enough, is present, and is derived/positive in the Bowman's low resolution .bam file.  However, one isolated downstream SNP is inconclusive.


The Homeland: In the footprints of the early Indo-Europeans.  Interactive map of ancient yDNA findings: link

Map of ancient yDNA at Indo-European.eu: link

Mr. I0806's belongings, above.