2 new Papers on ancient European DNA from UP to Neolithic

[Palermo Trapani]

Posth et al 2023 "Paleogenonomics of Upper Paleolithic to Neolithic European Hunter Gatherers"

https://www.nature.com/articles/s41586-023-05726-0

Villalba_Mouco et al 2023 " A 23,000 year old southern Iberian Individual links human groups that lived in Western Europe before and after the Last Glacial Maximum"

https://www.nature.com/articles/s41559-023-01987-0

If papers are already posted, my apologies and if the Moderators (Angela, Jovialis, Pax) think the 2 papers should be in 2 separate threads, I understand.

Cheers

 

[celtiberian-II]

The first paper is interesting:
"In conclusion, our study reveals that western and southwestern Europe served as climatic refugia for the persistence of human groups during the coldest phase of the last Ice Age whereas populations in the Italian peninsula and the eastern European plain were genetically overturned, challenging the role of these regions as glacial refugia for humans."

In the second paper:
"While more complex scenarios are possible, the observed genetic continuity suggests that the Iberian Peninsula, as a ‘southern genetic refugium’, could have sustained a stable population before, during and after the LGM, with no evidence for significant population turnover events but followed by an early and substantial contribution of Villabruna-like HG ancestry soon after."

 

[bicicleur 2]

still no clue about the origin of the WHG ..

 

[traveller]

“Our results confirm that part of the IUP ancestry present in Bacho Kiro and Tianyuan also survived in Southern Iberia MLZ until 23 cal kyr BP, ~12,000 yr later than the Aurignacian-associated Goyet Q116-1, the youngest previously known individual with traces of this ancestry. Initially, this IUP ancestry was attributed to East Asians as it is present in higher proportion in the Tianyuan individual, who shares more alleles with present-day East Asians than present-day or ancient Europeans26,45. The same type of ancestry was also observed in Goyet Q116-1 (ref. 26), who is more closely related to modern and ancient Europeans but still shares excess affinity to Tianyuan. Others45 have postulated an early pan-Eurasian population, which predated the split time of Europeans and Asians, as opposed to a back migration from Tianyuan-related Asian groups into Europe after the split. The oldest genomic data available from Bacho Kiro cave support the hypothesis of the existence of an Early Eurasian Bacho Kiro-like population that contributed to Ust’ Ishim, Tianyuan, Goyet Q116-1 and now also MLZ, but to the exclusion of other UP populations, including pre-Gravettian-associated Sunghir and Kostenki14, central European Gravettian or Magdalenian-associated individuals (Goyet Q2 cluster)”

 

[traveller]

“Here, we confirm a significant genetic affinity to Villabruna-like ancestry in pre-LGM Gravettian-associated individuals from central Europe but not (or to a much lesser extent) to Natufian-like ancestry. Of note, MLZ is one of the oldest individuals who shows a positive attraction to both Natufians (Z = 3.541) and Villabruna (Z = 2.972). By contrast, post-LGM populations share substantial ancestry with both Villabruna and Natufian but more with Villabruna than Natufians, in agreement with ref. 26 (Fig. 4a and Supplementary Table 2.10).

Others53 have shown that Natufians can be modelled using Villabruna-like ancestry and ‘Basal Eurasian’ ancestry, which constitutes an inferred population that diverged very early from all non-African populations after the split from African populations31. Our results show that MLZ shared an excess of Near Eastern ancestry present in Natufians, which is not explained by Villabruna-like ancestry itself (the oldest WHG with Near Eastern affinity). By contrast, we neither found an indication for Basal Eurasian ancestry in MLZ using other tests (Supplementary Information 6, Supplementary Fig. 6 and Supplementary Table 2.12) nor a high percentage of Neanderthal ancestry (Supplementary Information 7, Supplementary Fig. 7 and Supplementary Table 2.13).”

 

[bicicleur 2]

“Here, we confirm a significant genetic affinity to Villabruna-like ancestry in pre-LGM Gravettian-associated individuals from central Europe but not (or to a much lesser extent) to Natufian-like ancestry. Of note, MLZ is one of the oldest individuals who shows a positive attraction to both Natufians (Z = 3.541) and Villabruna (Z = 2.972). By contrast, post-LGM populations share substantial ancestry with both Villabruna and Natufian but more with Villabruna than Natufians, in agreement with ref. 26 (Fig. 4a and Supplementary Table 2.10).

Others53 have shown that Natufians can be modelled using Villabruna-like ancestry and ‘Basal Eurasian’ ancestry, which constitutes an inferred population that diverged very early from all non-African populations after the split from African populations31. Our results show that MLZ shared an excess of Near Eastern ancestry present in Natufians, which is not explained by Villabruna-like ancestry itself (the oldest WHG with Near Eastern affinity). By contrast, we neither found an indication for Basal Eurasian ancestry in MLZ using other tests (Supplementary Information 6, Supplementary Fig. 6 and Supplementary Table 2.12) nor a high percentage of Neanderthal ancestry (Supplementary Information 7, Supplementary Fig. 7 and Supplementary Table 2.13).”

View attachment 13779

Villabruna-like = Common west Eurasian ?
Natufian-like = Dzudzuana ?

 

[Palermo Trapani]

View attachment 13779

Villabruna-like = Common west Eurasian ?
Natufian-like = Dzudzuana ?

Kind of suggesting that Lazaridis and the team working on that Dzuduana paper that has been around since around 2018 I believe, which I don't think was ever published, might have been on to something. I don't have the paper in front of me but I seem to remember that paper suggesting Dzudzuana like ancestry (Near East/West Asia) present in populations to the West of the Caucuses very early on.

These results from Spain seem to have confirmed what Lazaradis et al back then were suggesting.

 

[Omino]

Kind of suggesting that Lazaridis and the team working on that Dzuduana paper that has been around since around 2018 I believe, which I don't think was ever published, might have been on to something. I don't have the paper in front of me but I seem to remember that paper suggesting Dzudzuana like ancestry (Near East/West Asia) present in populations to the West of the Caucuses very early on.

These results from Spain seem to have confirmed what Lazaradis et al back then were suggesting.

This one?
https://www.biorxiv.org/content/10.1101/423079v1

 

[bicicleur 2]

In qpAdm modeling, a deeply divergent hunter-gatherer lineage that contributed in relatively197 unmixed form to the much later hunter-gatherers of the Villabruna cluster is specified as
198 contributing to earlier hunter-gatherer groups (Gravettian Vestonice16: 35.7±11.3% and
199 Magdalenian ElMiron: 60.6±11.3%) and to populations of the Caucasus (Dzudzuana:
200 72.5±3.7%, virtually identical to that inferred using ADMIXTUREGRAPH). In Europe,
201 descendants of this lineage admixed with pre-existing hunter-gatherers related to Sunghir3
from Russia4 for the Gravettians and GoyetQ116-1 from Belgium3 202 for the Magdalenians,
while in the Near East it did so with Basal Eurasians. 203 Later Europeans prior to the arrival of
204 agriculture were the product of re-settlement of this lineage after ~15kya in mainland Europe,
205 while in eastern Europe they admixed with Siberian hunter-gatherers forming the WHG-ANE
206 cline of ancestry (Fig. 1c). In the Near East, the Dzudzuana-related population admixed with
207 North African-related ancestry in the Levant and with Siberian hunter-gatherer and eastern
208 non-African-related ancestry in Iran and the Caucasus. Thus, the highly differentiated
populations at the dawn of the Neolithic6 209 were primarily descended from Villabruna Cluster
210 and Dzudzuana-related ancestors, with varying degrees of additional input related to both
211 North Africa and Ancient North/East Eurasia whose proximate sources may be clarified by
212 future sampling of geographically and temporally intermediate populations.

 

[bicicleur 2]

Villabruna/WHG were the only descendants of this deeply divergent hunter-gatherer lineage that remained largely unadmixed.
They must have lived in an area that remains unsampled during the period 33-17 ka.
This area could be Ciskaukasia, the Ukraine or the Balkans.



Something like this maybe?

 

[Palermo Trapani]

This one?
https://www.biorxiv.org/content/10.1101/423079v1

Yes, that is the paper I was referring to. I haven't looked at it in a while but these 2 new papers made me think of that one. I need to go back and read it again.

 

[CrazyDonkey]

still no clue about the origin of the WHG ..

See the bolded/italicized text:

Fourth, we extend the finding of a large-scale genetic turnover as early as 14 ka in central and western European hunter-gatherers associated with multiple techno-complexes—Federmesser, Azilian and other Final Palaeolithic groups⁴—despite considerable technological continuity with the preceding late Magdalenian. This broadly distributed ancestry (the Oberkassel cluster (also known as WHG)) is most closely related to an Epigravettian-associated individual from northwestern Italy, suggesting that its expansion into continental Europe might have started from the west—and not the east—side of the Alps. Moreover, the almost complete genetic replacement of the Magdalenian-associated gene pool raises the hypothesis that parts of Europe were differentially populated during the abrupt climatic variation starting around 14.7 ka with the Bølling–Allerød warming period, creating areas where southern European populations could expand. This might also explain the genetic uniformity of the Oberkassel cluster across large parts of western Eurasia but genomic data from between 15 and 14 ka is needed to understand the exact dynamics of this turnover.

Fifth, the Oberkassel ancestry in western and central Europe and the Sidelkino ancestry in eastern Europe remained largely isolated for almost 6,000 years until genetic interactions were first observed—around 8 ka in northeastern Germany, possibly associated with cultural exchanges along the Baltics⁴⁹ and around 7.5 ka in the upper Volga region, possibly linked to the spread of pottery in the region⁵⁰.

In conclusion, our study reveals that western and southwestern Europe served as climatic refugia for the persistence of human groups during the coldest phase of the last Ice Age whereas populations in the Italian peninsula and the eastern European plain were genetically overturned, challenging the role of these regions as glacial refugia for humans. The incoming Villabruna ancestry later became the most widespread hunter-gatherer ancestry across Europe. Further palaeogenomic studies on Upper Palaeolithic individuals from the Balkans will be essential for understanding whether southeastern Europe represents the source of the Villabruna ancestry and a climatic refugium for human populations during the LGM.

 

[CrazyDonkey]

One previously unknown Gravettian lineage — dubbed Fournol, after a French site that is the earliest known location associated with this genetic cluster — inhabited what is now France and Spain. Another — named Věstonice after a Czech site — stretched across today's Czech Republic and Italy.

The Fournol descended from the Aurignacians, the earliest known hunter-gatherer culture in Europe, which lasted from about 43,000 to 33,000 years ago. In contrast, the Věstonice descended from the Kostenki and Sunghir groups farther east from what is now western Russia, who were contemporaries of the Aurignacians.

The Fournol genetic signature survived the Last Glacial Maximum, lasting for at least 20,000 years. Their descendants sought refuge in what is now Spain and southern France during the Last Glacial Maximum and later spread northeast to the rest of Europe.
In contrast, the Věstonice died out. Previously, scientists thought the Italian peninsula was a refuge for Gravettians during the Last Glacial Maximum, with the people there eventually forming the so-called Epigravettian culture after the glaciers retreated. However, the new findings show the Věstonice were not genetically detectable after the Last Glacial Maximum.

Instead, the new study finds the Epigravettians actually descended from Balkan groups that entered Italy as early as 17,000 years ago.

Starting about 14,000 years ago, the Epigravettians spread from the south across the rest of Europe, supplanting the Magdalenians, who were descended in part from the Fournol. The Magdalenians hunted reindeer that lived on the steppe, while the Epigravettians specialized in hunting forest prey. An abrupt warming event helped forests spread across Europe into what once was steppe, and the Epigravettians moved northward as well, Posth said.

https://www.livescience.com/unknown-lineage-of-ice-age-europeans-discovered-in-genetic-study

In other words, the Epigravettian culture was not the continuation of the Gravettian in the Balkins and Italy. (the Věstonice lineage), but their replacers. It was the Solutreans (the Fournol lineage), descended from the Aurignacians, who survived the LGM in Spain and France and then migrated into Europe, becoming the Magdalenian culture:

"Starting about 14,000 years ago, the Epigravettians spread from the south across the rest of Europe, supplanting the Magdalenians, who were descended in part from the Fournol."

This suggests that the supplanters of the Magdalenians could have been the Federmesser ("Quill Knife") culture ("late Upper Paleolithic to Mesolithic cultures of the Northern European Plain, dating to between 14,000 and 12,800 years ago" - https://en.wikipedia.org/wiki/Federmesser_culture) and related groups (Tjongerian, Creswellian, and Azilian cultures), eventually becoming the Ahrensburg culture c. 12,800 years ago, from which come the earliest finds of bows and arrows. They were superseded by the Maglemosian and Swiderian cultures.

 

[bicicleur 2]

https://www.livescience.com/unknown-lineage-of-ice-age-europeans-discovered-in-genetic-study

In other words, the Epigravettian culture was not the continuation of the Gravettian in the Balkins and Italy. (the Věstonice lineage), but their replacers. It was the Solutreans (the Fournol lineage), descended from the Aurignacians, who survived the LGM in Spain and France and then migrated into Europe, becoming the Magdalenian culture:

"Starting about 14,000 years ago, the Epigravettians spread from the south across the rest of Europe, supplanting the Magdalenians, who were descended in part from the Fournol."

This suggests that the supplanters of the Magdalenians could have been the Federmesser ("Quill Knife") culture ("late Upper Paleolithic to Mesolithic cultures of the Northern European Plain, dating to between 14,000 and 12,800 years ago" - https://en.wikipedia.org/wiki/Federmesser_culture) and related groups (Tjongerian, Creswellian, and Azilian cultures), eventually becoming the Ahrensburg culture c. 12,800 years ago, from which come the earliest finds of bows and arrows. They were superseded by the Maglemosian and Swiderian cultures.

the Magdalenians hunted mainly reindeer on the tundra with the atlatl
by 15.000 years ago they reached the melting icecaps in Denmark and northern Germany - the Hamburg culture

the first evidence of bow and arrow is at Stellmoor, northern Germany - the Ahrensburg culture, these were the people who replaced the Magdalenians in the north
so, not of all these WHG were forest hunters, as Posth claims

it looks like the Ahrensburg people didn't come from Italy, but directly from the Balkans, as you can see on this map




the SHG were a mixture of Iron Gate HG and EHG, they didn't have the late paleolithic Italian DNA

 

[CrazyDonkey]

it looks like the Ahrensburg people didn't come from Italy, but directly from the Balkans, as you can see on this map

I thought the language of the paper was confusing. If the "Epigravettians" replaced the Gravettian/Věstonice lineage in the Balkans and Italy from the east (or to the west), then coming from the Balkans would make the most sense, since they were blocked to the north by the Alps and to west by the Furnol. Italy would have been a cul-de-sac. When it says "the Epigravettians spread from the south across the rest of Europe," it doesn't say from where.

 

[MOESAN]

Aside the DNA question, it seems that even during the hardest times of LGM, hearths were found as far in North as Belgium, in Western Europe. Not as dense as in the Franco-Cantabric refugium of course. Maybe summer places for the most?

 

[jose luis]

Seasonal occupation during late spring/summer in Europe cold environments, even by Neanderthals and Protoaurignacians.


Subsistence of early anatomically modern humans in Europe as evidenced in the Protoaurignacian occupations of Fumane Cave, Italy
https://www.nature.com/articles/s41598-023-30059-3

Neanderthals in changing environments from MIS 5 to early MIS 4 in northern Central Europe – Integrating archaeological, (chrono)stratigraphic and paleoenvironmental evidence at the site of Lichtenberg
https://www.sciencedirect.com/science/article/abs/pii/S0277379122001500

 

[bicicleur 2]

Aside the DNA question, it seems that even during the hardest times of LGM, hearths were found as far in North as Belgium, in Western Europe. Not as dense as in the Franco-Cantabric refugium of course. Maybe summer places for the most?

maybe reindeer hunters following their prey up north in the summer?

 

[CrazyDonkey]

https://stoneageherbalist.substack.com/p/breakdown-two-new-genetics-papers

Magdalenian (17 - 12kya) - as the ice retreated, Solutrean derived groups moved out of Iberia and followed the vast reindeer herds into the continent. This culture produced much of the famous cave art.

Evidence for direct continuity between the Gravettian and the Magdalenian through the Solutrean period. The Solutrean individual Le Piage II from southern France has close links to the Fournol cluster and some affinity to the later Magdalenian cluster El Mirón. This indicates that the Solutreans arose from the western Gravettian people and maintained their population through the LGM.

All Epigravettians so far belong to the Villabruna cluster, and show no affinity to any Gravettian cluster. The oldest basal individual is the 13 kya Pradis 1 sample from north-east Italy. But given that Villabruna genetics appear at El Mirón some 19kya, the turnover must have been much earlier.

The Villabruna cluster shows a relationship with Near East hunter-gatherers, suggesting that they arrived in Italy from the Balkans.

Given that Villabruna affinities appear in both Iberia and the Balkans, it seems the LGM created a land and likely sea corridor from east to west, running underneath the glacial extremes.

The post-LGM Magdalenian culture has been characterised by both the El Mirón and GoyetQ2 clusters, suggesting that the Solutreans and Epigravettians mixed in some proportion to form the Magdalenians, which spread out north and north-east across Europe. New genomes confirm this, but El Mirón looks to be an outlier, with 43% Villabruna ancestry, whereas all other Magdalenian samples can be placed into the GoyetQ2 cluster.

Meanwhile a new group from the Balkans arrived in Italy and colonised the peninsula from north to south, as well as moving west to connect with the Solutreans. As the ice retreated, the Solutrean-Epigravettian mixture formed the Magdalenians, who swept northwards and east as far as Poland.

The WHG lineage can be better defined as the Oberkassel cluster, based on two 14kya newly sequenced individuals. This cluster shows the strongest affinity with a particular Epigravettian/Villabruna genome called Arene Candide16.

The Oberkassel cluster is made up of a large contribution from the Villabruna group and smaller amount from GoyetQ2 . All individuals within the WHG-Oberkassel group can be modelled as approx. 75% Arene Candide16 and 25% GoyetQ2.

Between 14-8kya, everyone in western and central Europe came from the Oberkassel lineage.

Possible additional Oberkassel ancestry was noted in samples from the upper Volga, but radiocarbon dating has not confirmed when this took place.

The main takeaways are: the Gravettians were not a homogenous population; the Epigravettians were an entirely separate population; the Solutreans maintained genetic continuity between the western Gravettians and Magdalenians; the Magdalenians were ultimately replaced by the WHGS; the WHGS and EHGs were largely isolated from one another until around 8kya.

Questions remain though about the relationship between the Epigravettians, the Near East hunter-gatherers, the Natufians and other pre and post LGM Eurasian populations. Now that we know the Epigravettians were descended from a different biological group, we need samples from those regions to unravel the formation of the Epigravettians, the Natufians, the Anatolian HGs and the Iberomaurusians.

The turnover from the Magdalenian to the WHGs seems extraordinary and the movements of different groups during the 8.2kya climate event need to be better understood to make sense of the EHG signal in western Europe - perhaps the introduction of pottery also came with a small number of exogamous marriages from east to west?

https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6478856/

In Europe, emerging ancient DNA evidence suggests that Late Glacial and Early Holocene population dynamics were associated with a major genetic substitution. This event saw the so-called Mirón genetic cluster dated c.19-15 kya and related to the Magdalenian culture replaced by the Villabruna genetic cluster at 14-7 kya¹⁹.

The population proxy shows five periods of significant positive, and four others of significant negative deviations from the exponential null model. The first of these occurs at the beginning of the time series, during the pleniglacial (18–17.6 kya), coinciding with the initial stages of the Middle Magdalenian culture. From 17.6 kya, we observe an abrupt decline followed by gradual increase in relative population densities. These changes occurred during the harsh climatic conditions of the Heinrich stadial 1 and the rapid warming of the Last Greenland Interstadial commencement (GI-1e or Bølling chronozone).

A second period of population increase is clearly identified between c.14-12.7 kya, showing minor fluctuations with three peaks (at 14–13.8, 13.7–13-4 and 13–12.7 kya) above the exponential null model of demographic growth, roughly covering the Final Upper Magdalenian-Early Azilian/Epipaleolithic transition. After the third brief spike (c.13–12.7 kya) the population densities abruptly decline from c.12.7-12.4 kya. In as far as we can be sure, given the natural limits of radiocarbon resolution which here is compounded in uncertainty by the marine reservoir effect²⁶, this decline coincides with the onset of the climatic effects of the Younger Dryas (YD) in the western Mediterranean. From 12.4 to 9.9 kya, we observe a long period of stable population sizes roughly covering the Epipaleolithic and the onset of the Early Mesolithic, manifesting in the data in three major negative deviations from the exponential null model at 11.7-11.3, 11.1-10.7 and 10.5–9.8 kya, covering most of the first half of the Early Holocene. This prolonged phase ended during the Early Mesolithic period, at c.9.8 kya, when the relative population densities abruptly increased to reach a positive pulse c.9.7–9.1 within the 95% CI envelope of the exponential null model. However, the high activity levels were not sustained; we observe next an abrupt decline in the population proxy, leading to a short period between 9 to 8.7 kya negatively departing from the null exponential model. Finally, we notice a pronounced recovery from 8.7 to 7.8 kya, during the Late Mesolithic. This last episode includes a narrow trough around 8 kya which might be correlated with the impact of the 8.2 kya cal BP cold event on Late Mesolithic populations^27,28. At the Iberian scale, however, this climatic event did not produce any statistically significant change to relative population levels. Finally, we observe a narrow peak in the population proxy at c. 7.8–7.59 overpassing the CI interval envelope of the exponential null model to abruptly fall again at the end of the time series.

he best fitting model of all was model F. The mean annual growth rates predicted by this model and their 95.4% CI are detailed in Table 1 and Fig. 5 (bottom). In this model, a distinctive demographic phase of exponential growth started with the Upper Magdalenian (c. 16.6 kya), (which covers different regional archaeological taxonomies, see Supplementary S⁷), and extends to the beginning of the Younger Dryas (c. 12.75 kya).

At this scale of analysis, constant changes to the environment could in theory have imposed a carrying capacity significantly less than the circumstances prevalent at the end of Demographic Phase 1. In light of this, we suggest that the abrupt decline in population levels c. 13 kya and the long period of stagnation of phase 2 could be explained by the effect of the abrupt environmental impact of the GS-1 stadial (YD) on foraging populations who had, at the end of the GI-1 interstadial, already reached (or were close to reaching) the carrying capacity of their environment. With the current data and for the patterns documented at the Iberian scale, a dynamic forager-resource disequilibrium model⁴⁷ could provide an explanatory framework to account for the population decrease and subsequent stationary demographic growth during the first half of the Early Holocene, especially changing levels of forest cover puts a limit on available resources.

These studies have suggested (i) a major population turnover at the end of the Late Glacial (c. 14.5 kya), with the reduction of post-LGM haplogroups (U2’3’4’7’8’9) and the subsequent major spread of haplogroup U5;⁷⁰ (ii) a major population replacement of the El Mirón aDNA genetic cluster (dated c.19-15 kya) by the so-called Villabruna cluster (dated c.14-7 kya) in Western Europe;¹⁹ and (iii) the admixture and persistence of populations with different relatedness with both genetic clusters in Iberia^71,72. In Western Europe, the ‘El Mirón Cluster’ consists of seven post-Last Glacial Maximum individuals from 19–14 kya, all associated with the Magdalenian culture. The Villabruna Cluster is composed of 15 post-Last Glacial Maximum individuals from across Europe dating between 14–7 kya years BP and associated with the Azilian, Epipaleolithic and Mesolithic cultures¹⁹.

Regarding the genetic replacement, it should be stressed that the date of the shift previously estimated in aDNA studies, 14 kya, derived from the date of the last known member of the El Mirón group, in Germany, and the first known member of the Villabruna group, a sample from Italy¹⁹. The paleogenomic evidence does not necessarily imply that the transition between the two genetic groups happened suddenly and simultaneously across Europe, and therefore must be addressed on a regional basis.

On the one hand, individuals with significant admixture proportions with the El Mirón cluster have been reported in Chan (NE Spain) at 9.1 kya, Carigüela (South Spain) at c.11.7–7.5 kya, Moita do Sebastião (Central Portugal) at 8.1 kya and Cingle del Mas Nou and Cocina Cave (E Spain) at 7.8 and 8 kya respectively^71,72. On the other hand, the first significant contribution of the Villabruna lineage is found at the Balma de Guilanyà (NE Iberia) at c. 12kya, and this group seemingly becomes dominant during the Late and Final Mesolithic in the Cantabrian region at La Braña c. 7.9 kya and Los Canes c. 7.1 kya⁷¹.

With the regional patterns documented in our study, the increasing population trend found in the Cantabrian subset from 8.5 kya onwards could be partially due to by the immigration of people associated with the Villabruna cluster.

 

[CrazyDonkey]

the Magdalenians hunted mainly reindeer on the tundra with the atlatl
by 15.000 years ago they reached the melting icecaps in Denmark and northern Germany - the Hamburg culture

the first evidence of bow and arrow is at Stellmoor, northern Germany - the Ahrensburg culture, these were the people who replaced the Magdalenians in the north so, not of all these WHG were forest hunters

There were population declines associated with sharp temperature declines:

1) From 17.6 kya, we observe an abrupt decline followed by gradual increase in relative population densities. These changes occurred during the harsh climatic conditions of the Heinrich stadial 1 and the rapid warming of the Last Greenland Interstadial commencement (GI-1e or Bølling chronozone).

2) "...the population densities abruptly decline from c.12.7-12.4 kya. In as far as we can be sure...this decline coincides with the onset of the climatic effects of the Younger Dryas (YD) in the western Mediterranean." The Younger Dryas' sharp temperature declines, following a sharp temperature increase, actually began ~14kya, not reversing itself until ~12.5kya. See: https://www.ncei.noaa.gov/sites/default/files/2021-11/3 The Younger Dryas -FINAL NOV (1).pdf.

https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6478856/

"The WHG lineage can be better defined as the Oberkassel cluster, based on two 14kya newly sequenced individuals. This cluster shows the strongest affinity with a particular Epigravettian/Villabruna genome called Arene Candide16....

The Oberkassel cluster is made up of a large contribution from the Villabruna group and smaller amount from GoyetQ2 . All individuals within the WHG-Oberkassel group can be modelled as approx. 75% Arene Candide16 and 25% GoyetQ2....

Between 14-8kya, everyone in western and central Europe came from the Oberkassel lineage." (Not from the GoyetQ2 lineage.)

"The post-LGM Magdalenian culture has been characterised by both the El Mirón and GoyetQ2 clusters, suggesting that the Solutreans and Epigravettians mixed in some proportion to form the Magdalenians, which spread out north and north-east across Europe. New genomes confirm this, but El Mirón looks to be an outlier, with 43% Villabruna ancestry, whereas all other Magdalenian samples can be placed into the GoyetQ2 cluster."

https://stoneageherbalist.substack.com/p/breakdown-two-new-genetics-papers

The Magdalenians were predominately GoyetQ2, but were replaced by those of Oberkassel/Villa Bruna ancestry, some of which could have come from El Miron. It could be that the Magdalenian "replacement" was largely a die-off due to harsh climatic reversals, with "forest dwellers" following the forest zone northwards, rather than directly competing with and replacing "rheindeer hunters" in the tundra.