Tag: G-BY211678 Haplogroup

This story is the third of a four part story focusing on the examination of possible social-cultural influences that may explain the lack of identified subclades (ancestors) in the migratory path of the Griffis genetic paternal line that lived in the Meuse Rhine watershed area. This third part of the story focuses on possible influences, notably patrilineal and matrilocal practices of in coming migratory groups, leading up to and during the bronze age.

The 2,850 year Gap between G-FGC7516 and G-Z6748: This phylogenetic gap was associated with haplogroup G-FGC7516. This common ancestor was born around 2200 BCE. The next genetic ancestor on the Griff(is)(es)(ith) YDNA line was associated with the genetic SNP mutation defining the G-Z6748 haplogroup, 2,850 years later. This gap represents about 95 generations.

There are a number of social, cultural and demographic factors that may have contributed to conditions that either limited the proliferation of G haplogroup subclades or were unfavorable for the preservation or detection of intermediate lineages over nearly three millennia for the Griff(is)(es)(ith) genetic line. The following factors can collectively explain why there is an absence of documented subclades during this time period:

• the convergence of early bronze age male bottlenecks;
• the impact of patrilineal and patrilocal social kinship structures in various migrating cultures that hindered G haplogroup proliferation;
• the long tradition of the Meuse Rhine watershed characterized as an area of social and cultural diversity; and
• border overlaps between various social groups along the Rhine River.

The Meuse-Rhine Water Basin as a Cultural and Geographical Crossroads Through Time

The Rhine-Meuse basin was not simply a transitional area, but an active frontier where contact, negotiation, and preservation of distinct cultural identities shaped the archaeological record and the evolution of traditions over time. Waterways facilitated connections, anchoring traditions and contacts both within the region and with distant areas. Despite episodes of external influence or migration, local adaptation and continuity remained central, producing a unique archaeological signature for the Rhine-Meuse basin over millennia.

In a research paper entitled “Archaeology from the Dutch Twilight Zone,” the archeologists David Fontijn, Patrick Valentijn, and Harry Fokkens explore the unique archaeological landscape of the Netherlands. They particularly focus on how local prehistoric communities shaped their environments and defined themselves in relation to wider European interaction networks during the Neolithic, Bronze Age, and Iron Ages.

The article delves into the Rhine-Meuse basin as a cultural meeting point for larger prehistoric networks, analyzing how social and cultural traditions intersected and changed over time. The archeologists introduce the concept of the “Dutch Twilight Zone” which captures the idea of the Netherlands as a liminal space, a cultural and geographic crossroads where different traditions and societies met and intermingled during prehistory (see illustration one). [1]

Illustration One: Looking at the Area of the Phylogenetic Gap in the Griff(is)(es)(ith) Genetic YDNA Lineage in the Context of Early Bronze Age Regional Achealogical Traditions

“(T)he Netherlands forms an interesting research area. On distribution maps it is generally placed outside or between large exchange and communication networks: the Nordic and the Atlantic spheres or traditions. Apparently the Netherlands constitutes a kind of marginal zone in between: a twilight zone. On the other hand the Netherlands, especially the Lower Rhine basin, is often referred to as a place of origin of developments, especially with respect to Bell Beakers. There is therefore a kind of paradoxical situation: it is in between, yet central.

“At the Rhine-Meuse delta two kinds of ‘borders’ seem to have coincided: a cultural border creating a social barrier and a traffic-geographical border creating a physical barrier. Neither was impenetrable, and indeed they were traversed in certain situations.” [2]

The archeologists substantiate the characterization of the Rhine-Meuse basin as a cultural meeting point for large prehistoric networks through several key archaeological observations:

Distinct Material Traditions: The authors note that while two large interaction networks (cultural and traffic-geographic) converged in the Rhine-Meuse region, major aspects of their respective traditions often remained separate. For example, metals from the southern ‘Atlantic’ sphere seldom entered the north, and ‘Nordic’ types rarely moved south. This points to a strong but structured interaction where contact did not necessarily lead to cultural blending.

Seafaring and Maritory Zones: The paper suggests that the strict division and nature of interactions were influenced by different coastal and tidal regimes north and south of the Rhine-Meuse area. Building on Stuart Needham’s concept of regional “maritories” (maritime territories) [3] , the researchers argue that frequent contacts within these zones were facilitated by coastal seafaring, fostering complex but localized exchange networks while retaining their respective cutlural patterns. (see illustration two).

illustration Two: Model of North Sea Maritories

Persistence of Local Cultural Traits: Archaeological evidence, such as the distribution of pottery types (e.g., biconical urns and Hilversum pottery) and burial traditions, demonstrates how local identities were maintained even as the area formed part of long-distance exchange systems. This persistence of distinct traditions alongside evidence for larger scale exchange and contact is taken as a hallmark of intersecting networks, “these regionally speciic archaeological cultures, were part of larger interregional exchange
networks“. [4]

Archaeological Findings and the Broader Region: Fokkens’ landscape-scale research documents how communities adapted to the region’s unique environment, and how influences from both the Atlantic and Nordic worlds can be discerned in settlement patterns, burial practices, and material culture.

“The Rhine-Meuse basin seems to have acted as a transition zone. Here we find evidence for exchanges between the two cultural traditions in the form of the occasional Funnel Beaker pot that occurs in Vlaardingen contexts, and Funnel Beaker knob-butted hammer axes in Stein contexts. However, the two networks do not seem to mix or amalgamate, they remain quite separate. There were no borders, yet cultural differences
are apparent.” [5]

Frontier and Border Zone: The Rhine-Meuse corridor, due to its geography, marked a natural boundary between different cultural worlds: the Atlantic to the southwest and the Nordic to the northeast. Evidence for this includes vessel forms, trade goods, and hoards that demonstrate both contact and separation. [6]

Selective Adoption and Exchange: Objects and practices such as Beaker pottery, metalwork, and funerary customs reveal selective adoption of external influences, combining local traditions with elements from wider European networks, rather than wholesale cultural assimilation.

“Against this cultural background around 2600 cal BC Bell Beakers appeared. First they occurred in Single Grave and Vlaardingen contexts, but after 2400 cal BC distinct regional groups developed. They seem to unify the existing cultural palette, but this is probably only supericial. The problem is that everyone focuses on burial contexts where Beakers seem to resemble each other in style, but if we look at settlement contexts, we can see that they were introduced in existing regional contexts. These regional traditions continued under a thin layer of Beaker ‘veneer’.” [7]

“(A)rchaeologists are far too ‘simple’ in their thinking about issues of social and cultural identity. No archaeologist today would speak of a monolithic, bounded archaeological culture reflecting ‘a people’. Nonetheless, in practice many still appear to maintain the notion that ethnicity has something to do with a totality of differences in ‘objective’ cultural traits. This is true despite an already established tradition of anthropological and sociological theory stressing that ethnicity is about situational self definition based on perceived cultural differences.” [8]

It is within this general context of a ‘Dutch twilight zone’ that perhaps we can understand why there are a number of factors that can explain the lack of identified subclades between haplogroups G-FGC7516 and G-Z6748.

“(T)he Dutch twilight zone appears to have been a transitional zone between two large interaction networks that ‘on the ground’ can be surprisingly easily distinguished; not only in regional cultural traditions, but also in the distribution of metal, house styles and burial traditions. The Rhine-Meuse delta appears to have been a border zone. It is as if people in the southern Netherlands felt part of a different world, travelled more frequently to the south and the west, and preferred to interact with the ‘Atlantic’ world. Over land the Meuse valley was probably one of the connecting routes, but coastal travel and cross channel travel probably was also ‘normal’. People from the northern uplands, on the other hand, looked more to the east and the north. Their burial and housing traditions were part of the Northern world and they probably ‘recognized’ more in the regionally different cultural traditions of the north than they did in the south.“ [9]

The Most Recent Common Ancestor Associated with the Beginning of the Phylogenetc Gap

For roughly 95 generations, the genetic YDNA ancestors of the Griff(is)(es)(ith) line lived and migrated within the ‘Dutch twilight zone’. The beginning of the phylogenetic gap starts with haplogroup G-FGC7516 (see illustration three). The genetic ancestor associated with this haplogroup had a 68 percent chance of being born between 2558 BCE and 1879 BCE. It is estimated that he was born around 2195 BCE which is around the advent of the bronze age.

Illustration Three: Scientific Details for haplogroup G-FGC7516

The Most Recent Common Ancestor of haplogroup G-FGC7516 lived in the general area of western Rhine River valley in what is currently known as the northern area of the Rhineland Palatinate of Germany (see illustration four). Around 2100 BCE, the primary cultural groups present in this area were societies that included early Celts and proto-Germanic peoples. Archaeological evidence from along the Rhine indicates significant settlement by these groups from 3000 to 1800 BCE, with the population engaging in farming, pottery, and early metallurgy during what archaeologists call the Neolithic and transitional Early Bronze Age.

Illustration Four: Location of MRCA of Haplogroup G-FGC7516 at the Beginning of the Phylogenetic Gap

Distinct Celtic and Germanic identities had not yet formed as seen in later history. Instead, there was a mosaic of local Neolithic farming communities connected to these larger continental traditions. 

The region maintained its importance due to its location along river trade routes and fertile land, facilitating a mix of local developments with influences arriving from the broader European context. During this period, the region was influenced by broader European prehistoric cultures, notably under the ‘thin cultural layer of veneer‘ from the Bell Beaker culture and the remnants of Corded Ware culture. These societies were ancestral to both Celtic and Germanic populations, but had not yet developed distinct Celtic or Germanic ethnic or linguistic identities by this time. [10]

Bronze Age Population Male Bottlenecks and Their Explanation

Patrilineal (father-to-son inheritance) and patrilocal or virolocal (women move to husband’s home) social systems leave distinct signatures on Y-chromosome diversity. Because men remain in their ancestral communities, dominant Y-chromosome lineages that are associated with dominant cultural groups become more homogeneous within local geograohical areas. Subordinate Y-chromosome lineages that are associated with minority cultural or social groups become less frequent. Meanwhile, incoming women from outside the dominant groups bring mitochondrial DNA diversity, but that does not affect the paternal lineages.

One of the most significant explanations for the lack of diversity and absence of Most Common Recent Ancestors (MCRAs) between haplogroups G-FGC7516 and G-Z6748 involves a well-documented genetic bottleneck that occurred during the transition from the Neolithic to the Bronze Age. This is tied to the the confluence of effects from the Corded Ware and Bell Beaker Cultures. Research has revealed a pronounced decline in male effective population sizes worldwide between roughly 3000 BCE and 1000 BCE that was not observed in female lineages. [11]

This genetic bottleneck would naturally result in the elimination of many Y-chromosome lineages, leaving fewer surviving male lines to develop subclades. This phenomenon, absent in female mitochondrial DNA, points to cultural and social dynamics rather than environmental factors. The bottleneck peaked at different times in the world. In Europe, it peaked around 3000 BCE or about 160 generations ago.

The bottleneck in Y-chromosome diversity coincided with significant cultural changes that affected the reproductive success among males. The decline in the male effective population size during this period was approximately one-twentieth of its original level in various regions of the world. This dramatic reduction would have eliminated numerous intermediate branches that might otherwise have existed in this time frame.

As mentioned in a prior story, two-thirds of all European men descend from just three haplogroup lineages identified as I1, R1a, and R1b in the late Neolithic Period. The descendants of the G2a haplogroup, such as the Griff(is)(es)(ith) YDAN lineage, experienced similar bottlenecks but did not experience as dramatic a subsequent proliferaion of lineages as did the I1, R1a, and R1 haplogroups in the bronze age. [12]

There are various explanations for this bottleneck. Earlier studies proposed that intergroup warfare between patrilineal clans caused repeated extinctions of male lineages. This hypothesis, supported by computer simulations, suggested that violent competition during the Neolithic/Bronze Age transition (when small-scale farming clans dominated) led to the loss of Y-chromosome diversity. [13] However, critics noted that even high rates of violence alone could not fully explain an observed 17:1 female-to-male effective population size ratio. [14]

Research linked the bottleneck to wealth accumulation during the Neolithic transition to agriculture. As societies shifted to agro-pastoralism, material resources enabled dominant males (and their sons) to monopolize reproductive success, drastically reducing genetic diversity among male lineages. [15] This “socially successful male” model aligns with archaeological evidence of emerging hierarchies but does not explicitly address the timing or severity of the bottleneck. [16]

A 2024 study by Léa Guyon and other researchers offers an alternative explanation centered on cultural evolution rather than violence. [17] The major factors associated with this hypothesis include:

• Patrilineal groups splitting into sub-clans of related males, homogenizing Y-chromosome lineages within groups;
• Expansions of some patrilineal groups and extinctions of others, amplified by patrilocal residence (male lineages staying localized while females migrated); and
• The bottleneck coincides with the spread of patrilineal inheritance systems followed by a recovery as bilateral kinship systems emerged.

This patrilineal system model replicates the genetic bottleneck using ethnographically documented non-violent social processes, showing that male effective population size could drop 20-fold within 100 generations through social dynamics alone.

Table One: Key Contrasts Between the Bronze Age Male Bottleneck Theories

Factor	Violence Hypothesis	Patrilineal System Model
Primary Driver	Warfare between clans	Cultural kinship rules
Male Mortality	High	Not required
Speed of Effect	Rapid diversity loss	Gradual, sustained decline
Supporting Evidence	Ancient conflict sites	Ancient DNA showing patrilocality & male-relatedness

While earlier theories emphasized conflict or inequality, the patrilineal systems hypothesis provides a parsimonious explanation that aligns with global genetic patterns and anthropological data associated with Neolithic social transitions.

G2a haplogroup lineages, initially common from earlier Neolithic populations, became marginalized as patrilineal and patrilocal social structures associated with the Corded Ware and Bell Beaker cultures favored rapid expansion of dominant male lineages (R1b) and reduced local Y-chromosome diversity for haplogroup G2a lineages. The decline of G2a subclades, such as the descendants of the most common recent ancestor of G-FGC7516, were the result of the successive generational effects of competition with dominant clans and marginalization and replacement of G2a clans (see table two).

Table Two: Mechanisms Influencing Haplogroup Decline through Successive Generations of Patrilineal and Patrilocal Descent

Social Influence	Description
Homogeneity within emerging dominant YDNA groups	Patrilineal descent meant G lineages became concentrated within specific clans and villages, reducing overall genetic variability but increasing homogeneity in each group. [18]
Female exogamy	Women migrated between groups, so mitochondrial DNA was mixed, but Y-DNA (such as G2a) remained fixed within patrilocal communities. [19]
Cultural hitchhiking and competition	Patrilineal societies caused certain haplogroups like G to either rapidly expand (if dominant) or be outcompeted. Intergroup competition or population bottlenecks could lead to extinction of entire G lineages if their male clans were wiped out or marginalized. [20]
Lineal fission	New clans or sub-clans often form by splitting along lines of paternal descent. This process clusters paternally related men together in the new group, compounding the concentration of specific Y-chromosome lineages and further restricting genetic diversity within each group. [21]
Patrilineal Power Dynamics	Social factors such as differences in social status among clans can promote the expansion of certain lineages if high-status men have more descendants, eventually dominating the group’s Y-chromosome pool and suppressing less prevalent lineages. [22]
Marginalization and replacement	Through demographic processes and male-biased migrations, G2a haplogroup’s frequency dropped as R1b and other steppe-derived lineages swept into the region and took over social structures, further exacerbated by patrilocal residence and patrilineal descent rules that favor the propagation of dominant male lines. [23]
Group extinction dynamics	If an entire clan or group becomes extinct (whether through competition, social dynamics, or other means), all the unique Y-chromosome lineages in that group are lost. Because clans are internally homogeneous for the Y chromosome, this means whole branches of Y-chromosome diversity can disappear, even if the overall male population size does not change dramatically. [24]

The Rhine River: The Corded Ware and Beaker Ware Cultures in the Muese-Rhine Watershed

The Rhine river formed an important boundary between two cultural zones, with the Corded Ware culture (c. 3100-2350 BCE) roughly to the east and the Bell Beaker culture (c. 2800-1800 BCE) spreading from the west and south. These cultures met at the Rhine, creating a dynamic contact zone that facilitated cultural exchange, technological innovation, and had effects on the composition of YDNA among the social groups. The Bell Beaker culture expanded eastward into Corded Ware territory from about 2400 BCE, marking a period of cultural contact and technological exchange. [25] Both of these cultures expanded throughout Europe (see illustration five).

Illustration Five: The Corded Ware and Bell Beaker Expansions

The Rhine contact zone acted as a demographic and cultural frontier where G haplogroups, once dominant among Neolithic farmers, faced steep decline due to Steppe-derived male-driven expansions. However, localized admixture in regions like the Rhine-Meuse delta allowed for fragmented survival of G lineages, particularly where pre-existing populations maintained ecological or social resilience. This dynamic underscores the interplay between migration, technological change, ecological barriers and regional adaptability in shaping genetic landscapes during the Neolithic-Bronze Age transition.

As indicated, the most recent common ancestor of haplogroup G-FGC7516 and his descendants lived during this time period in the vicinity of the middle region of the Rhine River. During this time period, the Meuse-Rhine watershed region was characterized by the interaction of the Corded Ware and Beaker Ware ‘archaeological cultures‘. The G2a haplogroup was representative of the YDNA admixture for both the Corded Ware and Beaker cultures (reference illustration six below).

Illustration Six: Corded Ware and Bell Beaker Cultures

“Archaeologists note that Corded Ware was not a “unified culture,” as Corded Ware groups inhabiting a vast geographical area from the Rhine to Volga seem to have regionally specific subsistence strategies and economies. There are differences in the material culture and in settlements and society. At the same time, they had several shared elements that are characteristic of all Corded Ware groups, such as their burial practices, pottery with “cord” decoration and unique stone-axes.” [26]

“The Bell Beaker phenomenon was not an ethnic culture like most other archeological cultures of the period, but rather represents a huge multicultural trade network inside which a variety of new artefacts, customs and ideas were exchanged and diffused, notably metalwork in copper, bronze and gold and archery.” [27]

As indicated in part two of this story, the Muese Rhine river watershed witnessed the limited adoption of Corded Ware pottery culture. It was introduced through haplogroup R1b-U106 (a steppe-associated lineage) to the region. The Funnelbeaker culture dominated much of north-central Europe, including the Rhine-Meuse region, from roughly 4300–2800 BCE. The Corded Ware culture appeared in the early third millennium BCE (around 2900–2650 BCE), eventually replacing the Funnelbeaker culture in this area.

Illustration Seven: Corded Ware Culture and the Griff(is)(es)(ith) Phylogenetic Gap

“We should observe that Corded Ware cultures co-existed for several hundred years with late Neolithic cultures … often residing in the close neighborhood, whether Globular Amphorae cultures in Poland or Bernburger cultures in southern Germany. What we observe in the archaeological record is, therefore, a gradual process of acculturation and integration which meant that, after 2400 BC, former strict cultural boundaries were being gradually dissolved and a new shared material culture appeared … . Bell Beaker groups had by now also emerged on the scene, introducing metallurgy and further complicating the mix of cultures and people. … (I)t took nearly a thousand years until all regions in northern and central Europe had adopted a shared social and cultural outlook that in all probability also included shared languages.” [28]

In the Meuse Rhine River watershed area, the adoption of Corded Ware pottery did not coincide with a significant influx of steppe ancestry. Individuals in these settlements exhibited Corded Ware Y-chromosome lineages but retained minimal steppe genetic input. This contrasts with other parts of Europe, where Corded Ware expansion brought substantial steppe ancestry. [29]

Illustration Eight: Distribution of the Bell Beaker Phenomenon

The Bell Beaker culture emerged later, around 2600–2500 BCE in the Lower Rhine area, arising through a fusion of local populations and Corded Ware-associated migrants (see table three). This new group then expanded rapidly, replacing or assimilating earlier Corded Ware groups and spreading across northwest Europe. The Bell Beaker culture is associated with its distinctive bell-shaped pottery, copper daggers, and new burial customs. [30]

Table three provides a rough correlation between the Griff(is)(es)(ith) YDNA lineage with the population events in the Meuse- Rhine Region. The ancestors that were associated haplogroups that lived earlier than 2500 BCE may have lived in an area southwest of the Meuse Rhine area.

Table Three: Key Population Events in the Rhine-Meuse Region

Time Period	Griff(is)(es)(ith Haplogroup	Major Event	Genetic Impact
6500–4000 BCE	G-L497, G-CTS9737, G-Z1900, G-Z690	Early Neolithic farmer expansion	Limited in Rhine-Meuse; hunter-gatherer continuity along with Neolithic farming groups
~3000–2500 BCE	G-Z1817, G-Z727	Corded Ware complex arrives (incorporating Neolithic groups)	Little steppe ancestry; Corded Ware Y-chromosome present
~2500 BCE	G-Z1817, G-Z727, G-FGC477	Formation of Bell Beaker groups (incorporating local & Corded Ware groups)	Major admixture, new expansive population
Post-2500 BCE	G-FGC7516	Bell Beaker expansion	Disruption of local continuity, major population replacement in northwest Europe

Over successive generations, the interaction between the Corded Ware and Bell Beaker cultures along the Rhine during the Late Neolithic and Early Bronze Age (2200–1800 BCE) likely had an influence on impeding the growth and distribution of G2a haplogroup lineages through patrilineal and patrilocal customs of the Corded Ware and Bell Beaker groups. In marginal or ecologically distinct zones (e.g., wetlands of the Rhine-Meuse delta), local populations with members of the haplogroup G2a may have resisted full assimilation, as seen in the prolonged coexistence of Funnelbeaker and Corded Ware groups. [31] This could explain sporadic G2a haplogroup survival, such as the Griff(is)(es)(ith) paternal line, in areas less dominated by Steppe migrations.

A study by Ralph Großmann underlines that the relationships between Corded Ware and Bell Beaker phenomena were complex, involving shared practices and hybrid identities rather than sharp cultural divisions. While the Corded Ware and Bell Beaker complexes of the third millennium BCE are often seen as distinct in terms of burial rituals and material culture, there are significant overlaps and interrelations between them, particularly in regions where their cultural spheres were geographically adjacent. [31a]

In regions like the Rhine and Thuringia Basins, burial practices and vessel decorations from the two cultures display notable similarities and overlap, especially where communities interacted closely in space and time. Overlaps are most pronounced near river confluences, suggesting that these were hubs of cultural interaction and exchange. The evidence points to more fluid and negotiable social identities in these regions, with possible hybrid or multiple cultural affiliations rather than strictly bounded group identities. [31b]

The Corded Ware Culture: Effects of a Patriachical Society

“For the Corded Ware complex it has been suggested that initial migrations were dominated by males, who married in women probably from residing Neolithic populations, although at present it is debated whether the genetic evidence for male-dominated migrations contributing to these groups is compelling. There is also evidence that the individual groups most likely practiced patrilocality and exogamy at a community level.” [32]

An innovative research paper by Kristia Kristiansen and colleagues integrate recent breakthroughs in genetics, stable isotope analysis, archaeology, and linguistics to re-examine how the Corded Ware Culture formed in Europe after 3000 BCE. They have documented that migration was central to the formation of the Corded Ware Culture, with ancient DNA evidence confirming a large-scale movement of Yamnaya pastoralists from the Pontic-Caspian steppe into Europe after 3000 BC. These migrations did not result in mere population replacement. The formation of Corded Ware societies was driven by complex local interactions between the incoming Yamnaya and indigenous Neolithic groups. [37]

Corded Ware societies emerged through the blending of Yamnaya migrants and local Neolithic populations. This process involved not just biological – genetic mixing, but also the exchange of cultural practices, farming techniques, and language. [38]

The integration between the migratory groups and the indigenous Neolithic groups was facilitated by exogamic marriages, particularly unions between Yamnaya men and Neolithic women. This role or mechanism of exogamic marriage created a social foundation for new dialects and material cultural forms as pottery skills and linguistic influences were exchanged. It also had a diminishing effect on Neolithic YDNA G2a subclades such as the Griff(is)(es)(ith) paternal line. Illustration nine depicts the model of the social processes associated with exogamy that transformed Yamnaya culture to Corded Ware culture (labled “A”) and its subsequent migratory effects as Corded Ware culture lead to further adaptation and transformation (labeled “B”).

Illustration Nine: Social Processes of Exogamy Transforming Yamnaya to Corded Ware Culture and Its Subsequent Migration

An article by Quentin Bourgeois and Erik Kroon argues that a Corded Ware identity reflects the sharing of world views, practices, and ideas—not just common genetic backgrounds. Burial practices serve as a crucial lens for understanding the cultural transmission and social networks in Corded Ware communities. Their analysis of 1,161 Corded Ware burials across north-western Europe shows that men’s burials were much more similar across vast distances and regions, whereas women’s burial practices remained distinctly local. [39]

Illustration Ten: Distribution map of the Corded Ware burials in Bourgeois & Koon Study

Male burials featured standard positions (semi-flexed on the right side, head west and facing south) and similar grave goods, whereas female burials had more local variations. The striking similarity in male burials indicates that male burial ritual was the main vector for transmitting Corded Ware identity and funeral customs across different communities. This international character of male burials suggests that men played a central role in maintaining and spreading shared cultural practices. [40]

The differences and similarities in burial rites point to a complex society where the role of gender was significant. The uniformity in male burials, despite regional diversity, supports the theory that the Corded Ware society was male-focused, aligning with recent ancient DNA studies showing male-dominated migration patterns. [41]

Migration introduced new forms of social organization, reflected in burial customs and property transmission (individual rather than collective), with Corded Ware dominance surfacing in the centuries leading into the Bronze Age. While there was cooperation and intermarriage, evidence also indicates episodes of conflict and local diversity in the outcomes, with variation in material culture and genes depending on region. [42]

The Corded Ware culture is widely regarded as a major vector for the spread of Indo-European languages in Europe, particularly Proto-Germanic and Proto-Balto-Slavic. Its social and burial practices influenced subsequent cultures, such as the Bell Beaker culture, and contributed to the development of individualized chiefdoms and early Bronze Age societies.

An article by Kroon and other researchers highlights that the introduction of the Corded Ware Culture during the Late Neolithic on the Dutch West Coast was not a uniform process, but rather varied significantly from site to site. A core finding is the remarkable continuity of ceramic technological traditions across the region, suggesting that social practices and local technological knowledge persisted even as new material styles were introduced. [43]

The study used ceramics as a proxy for understanding cultural and social changes, applying geochemical, mineralogical, and morphological analyses to pottery from three Late Neolithic sites in the Dutch coastal area. The authors compared three hypotheses for how Corded Ware culture might have spread: migration, diffusion, and local network interactions.

There is strong evidence for continuity in ceramic production techniques across the transition to Corded Ware culture, indicating local populations maintained their methods despite adopting new styles. The adoption of Corded Ware material culture reflected a local process of cultural integration, rather than a wholesale replacement or abrupt change. The impact of Corded Ware culture introduction was site-dependent, varying in degree and manner across the studied locations. [44]

The results challenge views that see the Corded Ware phenomenon solely as a result of migration or large-scale population replacement. The study demonstrates that archaeological changes in material culture (such as pottery types) may not always indicate deep social rupture, but might instead reflect complex and regionally specific adaptations. These insights from the study emphasize the relevance of technological traditions as markers of social continuity during periods of apparent cultural change in prehistory.

Table Four: Summary of Key Features of Corded Ware Culture [45]

Feature	Description
Pottery	Beakers and amphorae (e.g. jars with two handles) with corded decoration
Burial	Single inhumations under barrows; Gendered body orientation: Males were typically placed on their right side with their heads positioned towards the west; Females were typically placed on their left side with their heads positioned towards the east; and Both genders, regardless of their orientation, faced towards the south.
Weapons	Stone battle axes, flint arrowheads, some copper items
Economy	Primitive agriculture, animal husbandry, possible horse domestication and likely with an emphasis on cattle and sheep.
Social Structure	Individualized, patriarchal, emerging social hierarchy: the society appears to have been stratified, with increasing social differentiation and competition for prestige goods and raw materials such as flint, stone, and amber.
Land Use	Depending on region, may have lived in villages, hamlets, and farmsteads, with houses that were usually small and divided into two or three spaces.
Mobility	Use of wheeled vehicles; evidence of long-distance trade and semi-nomadic pastoralism
Regional Diversity	Multiple local variants; influenced by earlier and neighboring cultures. It displayed significant regional variation in pottery styles and burial customs, leading to the identification of several regional groups (e.g., Single Grave, Battle Axe, Fatyanovo cultures).
Kinship Patterns	Archaeological and burial evidence points to a patriarchal society, with male burials often more elaborate and associated with weapons, suggesting the prominence of warrior elites.
Language Impact	Spread of Indo-European languages: associated with the diffusion of Proto-Germanic and Proto-Balto-Slavic speakers, and some believe that the people of the Corded Ware Culture spoke a steppe language that contributed to some of the contemporary Indo-European languages.

The Bell Beaker Culture: Continued Impact of Patrilineal and Matrilocal Cultural Social Practices

Before the arrival of Bell Beaker groups, the Corded Ware culture—descendants of steppe populations with significant Yamnaya ancestry—became established in the Netherlands and across northern Europe. These people brought distinctive genetic markers (such as Y-chromosome haplogroup R1b-M269) into the region. When Bell Beaker practices spread, they often did so among populations already rich in steppe-related ancestry, especially in the central and northern Netherlands. [46]

Around 2500 BCE, a significant demographic shift occurred with the emergence of Bell Beaker-associated populations. These groups arose through the fusion of local Rhine-Meuse inhabitants (contributing 9–17% of ancestry) and Corded Ware-associated migrants of both sexes. This new population expanded rapidly from the Rhine-Meuse region, disrupting the long-standing local continuity. [47]

The interaction between the dominant Corded Ware and Bell Beaker cultures along the Rhine during the Late Neolithic and Early Bronze Age (2200–1800 BCE) likely had an influence on limiting the growth and distribution and G haplogroup lineages, particularly the descendants of the most common recent ancestor of G-FGC7516.

Anthropometric analysis of the skeletons enabled the determination of sex and age at death. Strontium and oxygen isotope ratios in tooth enamel were determined to find out if individuals grew up at similar or different geographical locations. This, combined with the sequence analysis of DNA fragments from bones, provided evidence for a patrilocal society in both communities. Whereas males shared a single Y-haplotype there was a much higher maternal genetic diversity, with 23 mitochondrial haplotypes in 34 individuals.

The reconstructed pedigrees were consistent with monogamy combined with exogamy, as six of eight non-locals were women. successive generations of males remained in these communities and women came from elsewhere. In this way property such as farming grounds could be conserved and inbreeding was prevented. The single Y-haplotype, with a Steppe-pastoralist origin from Eurasia, is still the dominant type in central and western Europe today.

“By combining the various sciences, and applying them to the 42 graves of our two late Bell Beaker culture cemeteries, (the researchers proposed) a model characterized by six social principles:

1. The basic kinship units are nuclear families. By this we simply imply a small family group rather than an extended family group. ...
2. These nuclear family groups are based on patriarchal, patrilinear and patrilocal residency
   lines. …
3. The marriage system is based on female exogamy and likely monogamous. …
4. The inheritance system is likely based on male primogeniture. …
5. Nuclear families likely formed independent households. … (and)
6. Families/households formed alliances through kinship and the observed exogamic practices and foster children further forged such alliances, likely linking families into clans. Alliances were thus regional rather than closely local, and they could have formed larger political and ethnic entities to be mobilized in periods of unrest, or during periods of expansion.” [49]

Illustration eleven provides a schematic depiction of how Bell Beaker patrilineal based new settlements were established and consolidated through the marriage of Corded Ware men and Neolithic females from outside groups. In the context of the expansionist kinship system revealed by the study, the concept of foster sons also plays a significant role, both in the Bell Beaker communities discussed in their study and in later Indo-European societies. Foster sons acted as living links between families and communities, cementing the cooperative and expansionist orientation of Bell Beaker society.

Illustration Eleven: Bell Beaker Kinship and Social Organization

The study notes that such patterns—male-centered descent, the preferential burial of young boys, and exogamous marriage patterns—reflect kinship mechanisms that later appear in Indo-European societies, where fosterage of sons becomes a well-attested tradition (see table five).

Table Five: Role and Utilization of Foster Sons in Bell Beaker Culture [50]

Role	Utilization
Fosterage as Alliance-Building	Foster sons (“fosterage”) involved sending young boys to be raised by families outside their birth group, often with allied families or communities. This practice built social and political ties, extended trust networks, and reinforced alliances between distinct lineages or groups.
Socialization and Integration	Foster sons would become closely affiliated with their host families, forming almost familial bonds with their non-biological “parents” and siblings. This system allowed boys to gain social capital, language skills, and knowledge of other communities, making them valuable cultural and diplomatic bridges.
Expansionist Strategy	By fostering sons out to other communities, Bell Beaker groups could strengthen inter-community relationships, reduce the risk of conflict, and facilitate trade and cooperation. This practice also helped to integrate and stabilize rapidly expanding social networks as these communities spread across Europe.
Elite Status and Selectivity	The archaeological bias for burying male youths may be a reflection of this practice. Communities invested more in the status and commemoration of boys, especially those with ties (by blood or fosterage) that reinforced the kin-group’s reach and alliances.

The study by Karl-Göran Sjögren, Volker Heyd and other researchers provide cogent aguments based on empirical research regarding the influence of Bell Beaker kinship social patterns on migration and admixture of YDNA haplogroups. Their study focused on Bell Beaker settlements in what are close contemporary areas of lower Bavaria Germany (see illustration twelve). These settlements were southwest of the Meusse Rhine River watershed area where Griff(is)(es)(ith) YDNA descendants lived during the time period that is being discussed in this story.

Illustration Twelve: Location of Two Bell Beaker Culture Cemetery Sites

One might question the validity of generalizing the conclusions of this and other related studies for their apllicability to the Muese Rhine Watershed area. As stated, the Bell Beaker culture represented a huge multicultural trade network with many local regional variations. It is possible and probable, despite the distance of roughly 420 miles between the two geographical areas, that the socio-cultural characterteristics may have been very similar between the two geographic areas. [51]

“A wide range of regional diversity persists within the widespread late Beaker culture, particularly in local burial styles (including incidences of cremation rather than burial), housing styles, economic profile, and local ceramic wares (Begleitkeramik). Nonetheless, according to Lemercier (2018) the mature phase of the Beaker culture represents ‘the appearance of a kind of Bell Beaker civilization of continental scale’.“ [52]

The Bell Beaker culture of the Meuse-Rhine region was dynamic and characterized by social hierarchies, diverse burial and material practices, long-distance exchange, and interaction with neighboring groups, all within a context of local adaptation. [53]

The use of shared burials for adults and children likely had symbolic meaning, possibly highlighting family status, lineage, or emphasizing transgenerational links within communities. These practices also mark the transition from collective to individual burials, underlining a shift in attitudes toward family, children, and social identity. [56]

The inclusion of substitute parents (such as a paternal aunt) in burials indicates that extended family members took on caregiver roles, sometimes substituting for biological parents in both social life and burial practices. This suggests that Beaker social systems were flexible and valued non-nuclear familial bonds, especially on the paternal side. [57]

The similarities in grave goods, burial postures, and arrangement across geographically distant sites (e.g., Luxembourg and Britain) point to widely held, highly formalized burial rituals. These reflect community-wide standards of mourning and memorializing, indicating coordinated cultural beliefs and behaviors across Beaker regions—independent of local variation. [58]

Regional Variations of the Beaker Culture in the Meuse Rhine Watershed Area

The genetic profiles of Dutch Bell Beaker individuals show admixture not only from incoming steppe-related groups but also from earlier European Neolithic farmers and indigenous hunter-gatherers. This created a heterogeneous genetic makeup, visible in the variety of mitochondrial and Y-chromosome haplogroups found in Beaker-era remains, and evidenced by studies showing a mix of ancestries within local sites. [59]

Regional backgrounds played a crucial role in shaping the development of the Bell Beaker culture in the Meuse Rhine watershed area. Instead of a single, straightforward evolution, the transition to Bell Beaker culture occurred differently across various Dutch regions, reflecting their unique cultural and settlement histories. [60]

The Netherlands sat at a crossroad between different Late Neolithic cultures: the Single Grave Culture (a variant of the Corded Ware), the Vlaardingen Culture to the west, and continued contact with regions to the south and east. This location fostered the blending of traditions—forms, decorated pottery, and burial customs—resulting in Dutch Bell Beaker culture that sometimes bridged or combined elements from neighboring regions rather than simply copying them. [61]

Where some Dutch regions, particularly in the central and east, show continuity from Single Grave to Bell Beaker traditions, the west and south (with more influence from the Vlaardingen culture) witnessed abrupt transitions as external Beaker elements were introduced and reshaped local traditions. These varied pathways demonstrate that cross-cultural interaction, involving migration, exchange, and local adaptation, was central to Beaker diversity in the Meuse Rhine watershed area. [62]

Cross-cultural interactions, including the movement of clans, intermarriage, trade, and the selective adoption of foreign elements, fueled the diversity of the Beaker phenomenon in the Netherlands, with each region reflecting the unique ways that local and external traditions blended. [63]

Beakers, more than any artifact series in Dutch prehistory, are “international” in character, reflecting the Netherlands’ position in broader European trade and communication networks. The multiplicity of Beaker forms and practices found locally document these wide-ranging cultural influences. [64]

Social dynamics were shaped by competition among descent groups and individuals for prestige, status, power, and wealth. This competition fostered a moderate degree of social ranking and the emergence of local elites, but did not reach the full social stratification seen in later chiefdoms or state structures. Leadership was likely in the hands of local elites who controlled the production and distribution of valuable goods, such as the characteristic bell-shaped pottery. Personal prestige and status were important, as reflected in burial practices and grave goods.

A study by Marc Vander Linden challenges the traditional interpretation of third millennium BCE European societies as being dominated by rigid social hierarchies, especially in the context of the Bell Beaker culture. Rather than societies strictly organized around elites competing for prestige, Vander Linden’s research suggests that the Bell Beaker period saw the emergence of more fluid and dynamic social structures. These were marked by increased mobility and the widespread circulation of people, ideas, and material culture. The evidence from southern France indicates that the Bell Beaker culture fostered integration and connectivity between different groups, promoting collaboration and the formation of wider social networks. This perspective puts less emphasis on the accumulation and display of wealth by a small group of elites. [64a]

The study urges archaeologists to reconsider the evidence for social hierarchy and the narrative of bronze-age societies progressing linearly toward greater stratification. Instead, the Bell Beaker phenomenon may represent an alternative model of community building and interaction, centered on shared practices and social cohesion. Vander Linden’s interpretation broadens the understanding of prehistoric European social organization, highlighting the role of networks, exchange, and integration rather than just competition and hierarchy. This can impact how scholars interpret similar patterns in other regions and periods, questioning the assumed inevitability of hierarchical development in early complex societies.

As summarized in table six, the Bell Beaker culture in the Meuse-Rhine watershed area is characterized by several key features of social organization and culture, as revealed by archaeological and recent scientific research.

Table Six: Notable Features of Bell Beaker Culture in the Meuse Rhine Watershed Area

Feature	Description
Hybrid Identities and Cultural Contact	The Meuse-Rhine watershed was a region where different groups met and interacted, especially at river confluences. This led to overlapping characteristics in burial and material culture, such as vessel decoration styles shared between Bell Beaker and Corded Ware groups. The result was a fluid social identity and occasional hybridization. [65]
Trade and Mobility	While many groups practiced sedentary farming, others maintained mobile lifestyles, possibly as traders or pastoralists. This diverse settlement pattern likely contributed to the rapid spread of Bell Beaker goods and culture across Europe. The Bell Beaker people were part of extensive exchange networks, moving raw materials such as copper over long distances. The Rhine-Meuse area served as an essential network node, facilitating interactions between different cultural groups, and the spread of both population and ideas. Bell Beaker people relied on and innovated sea-based and coastal routes to expand their culture widely across Europe, with a particular focus on the Atlantic littoral and the Mediterranean basin. [66]
Settlement Patterns & Landscape	Bell Beaker groups typically lived in small settlements, rather than large, centralized societies. Typical Bell Beaker settlements housed small villages of 30 to 50 inhabitants. Archaeological evidence suggests that the Bell Beaker people inhabited both sedentary farming settlements and maintained mobile elements. The settlement structure in the region adapted to the local landscape—such as river valleys and higher ground, mirroring an ability to blend new ideas with existing traditions. [67]
Kinship Patterns	Social life was organized around descent groups, with kinship (biological or mythical) being the main organizing principle. These groups likely had a clan structure, often exogamous and patrilineal, emphasizing descent from a common ancestor. [68]
Social Structure	The presence of specialized craft goods suggests an economy that valued certain occupations, such as metalworking. Some settlements likely functioned as specialized production or trade centers for particular goods. Emerging social hierarchy: the society appears to have been stratified, with increasing social differentiation and competition for prestige goods and raw materials such as flint, stone, and amber. [69]
Burial	Bell Beaker burial styles included single, crouched inhumations in individual graves, often with a bell-shaped pot, flint arrowheads, and a stone wrist guard as grave goods, and typically located on well-drained soil. Burial orientation varied by region and sex, with males in Continental Europe generally on their left side and females on their right, both facing east. Graves could be flat or covered by circular mounds, with some evidence suggesting extended family burials and a focus on kin relationships.  [70]
Burial & Symbolic Practices	Not every person received a Bell Beaker burial, which suggests that burial with bell beaker ceramics and goods was a selective, perhaps prestigious, rite reserved for particular individuals or events. [71]
Pottery	Regional diversity of Bell Beaker ceramics in the northwestern Mediterranean (Spain, France, Italy). This diversity, evident in decorative styles and burial contexts, enables researchers to construct detailed chronological sequences and to trace the arrival, adoption, and adaptation of Bell Beaker elements within local societies. [72]

Challenges of Poor Skeletal Preservation in the Meuse Rhine Watershed Area

The Meuse-Rhine watershed is not an ideal environment for long-term skeletal preservation due to its high humidity and waterlogged conditions. DNA is vulnerable to degradation from microbial action, hydrolysis, and oxidation, all of which are exacerbated in damp environments. Many archaeological sites in the region contain only small, morphologically unidentifiable bone fragments, rather than complete skeletons. Environmental factors like soil properties, temperature, and exposure to humidity are significant factors in how well bone and DNA are preserved. [73]

Despite widespread poor skeletal preservation in the Meuse-Rhine watershed due to adverse environmental conditions, significant ancient DNA (aDNA) data have been successfully recovered. Researchers have overcome the challenges of poor preservation by targeting denser bone elements like the petrous bone and teeth, where DNA is better protected. The region’s archaeological record, though subject to decay from its riverine and wetland environments, has yielded substantial genetic information that has reshaped the understanding of European prehistory. [74]

Archaeologists and paleogeneticists have successfully extracted extensive aDNA from the Meuse-Rhine area, demonstrating that usable genetic material is available even in poorly preserved skeletal remains. The dense petrous part of the temporal bone and the cementum of teeth are known to protect DNA from environmental decay far better than other skeletal elements. For example, a 2025 study was able to assemble genome-wide data for 109 people dating from 8500–1700 BCE from the Rhine-Meuse area. The success of this study in collecting a large dataset confirms that despite poor overall preservation, high-yield specimens can be found. [75]

The ancient DNA data recovered from the Meuse-Rhine area have provided groundbreaking insights into the region’s population history, particularly for the periods of 8500–1700 BCE. Genetic analysis revealed a high proportion of hunter-gatherer ancestry persisting in the Rhine-Meuse area for three millennia longer than in many other parts of continental Europe. The data document a unique demographic transition. Unlike other regions that saw a near-complete turnover with the arrival of farmers, the Rhine-Meuse area experienced persistent hunter-gatherer presence and an exceptional pattern of interaction with incoming populations.

The genetic analysis also showed that the arrival of the Bell Beaker complex in the western Netherlands was unique. The local lowland population that adopted the pottery had very little steppe ancestry, unlike their counterparts elsewhere in Europe. [76]

“The spread of CW (Corded Ware) influence to the wider Rhine-Meuse area was more complex than in many areas of central and eastern Europe. In the uplands, where skeletal material tends to be poorly preserved and no ancient DNA data are available, the complete CW package emerged; marked by the construction of CW burial mounds, the general absence of settlements, and sparse pottery finds.” [77]

“The arrival of the BB (Bell Beaker) complex around 2500 BCE marked another major cultural transition, as settlements spread across the wetlands and coastal areas, replacing Vlaardingen/CW settlements, though generally not using the same sites. The BB economy was similar to the previous CW one and consisted of predominantly farming mixed with low-intensity hunting and gathering. In the sandy uplands, there was a continuation of the barrow ritual, but with distinct BB characteristics and material culture replacing the CW repertoire. BB groups were also well attested south of the Rhine, as evident in BB burial mounds on the sandy soils of the southern Netherlands and Belgium. BB settlement sites remain just as elusive in this area as CW settlements. However, the presence of ploughland dated to the Late Neolithic suggests that the lack of settlement evidence is not the result of nomadism but rather of settlements in lower lying places where there is little chance for detection by archaeologists.“ [78]

While there have been technological inovations in extracting aDNA from poorly preserved remains, a challenge still exists in finding skeletal remains of individuals who represent YDNA haplogroups, such as various subclades in the G2a hapklogroup, that have been marginalized through time.

The Next Part of this Story

The fourth part of this story discusses the enduring social, cultural and demographic factors that may have contributed to the continued lack of subclade development for the Griff(is)(es)(ith) paternal YDNA line during and after the bronze age up to the end of the phylogenic gap around 650 CE.

The lack of identified YDNA subclades documenting the migratory path in the Meuse Rhine watershed area can be partly attributed to three major demographic factors that limited the subsequent growth of G2a subclades, particularly the Griff(is)(es)ith) paternal line:

1. the persistence and admixture of I2 and C1 hunter-gatherer haplogroups with G2a farming haplogroups in the late neolithic;
2. the enduring impact of R1b Bell Beaker migrating groups into the area in the post Neolithic and early Bronze age; and
3. the continued dominance of R1b social and cultural factors on G2a subclade growth.

Source:

Feature Banner: The banner at the top of the story features a map of the phylogenetic gaps discussed in the story. The map was generated by taking a snapshop from the FamilyTreeDNA Globetrekker^TM video of the migratory path of my YDNA descendants over time. The map shows the migratory path of selected most common recent ancestors and their respective estimated dates of birth. The table in the middle of the banner points to the influence of the Corded Ware and Bell Beaker cultures. The illustration on the right provides a schematic depiction of how Bell Beaker patrilineal based new settlements were established and consolidated through the marriage of Corded Ware men and Neolithic females from outside groups. This social process had a negative impact on the proliferation of G2a YDNA subclades.

[1] David Fontijn, Patrick Valentijn, and Harry Fokkens, Archaeology from the Dutch Twilight Zone, in Sophie Bergerbrant and Serena Sabatini (ed), Counterpoint: Essays in Archaeology and Heritage Studies in Honour of Professor Kristian Kristiansen, Oxford: Archaeopress, 2013, 531-540 , https://www.academia.edu/6072124/Archaeology_of_the_Dutch_Twilight_Zone

[2] David Fontijn, Patrick Valentijn, and Harry Fokkens, Archaeology from the Dutch Twilight Zone, Page 539

[3] Needham, Stuart, Encompassing the Sea: ‘Maritories’, and Bronze Age maritime interactions, 12-37, in Peter Clark, ed, Bronze Age Connections Cultural Contact in Prehistoric Europe, Oxford: Oxbow Books, 2006 https://dokumen.pub/qdownload/bronze-age-connections-cultural-contact-in-prehistoric-europe-1842173480-9781842173480.html

[4] David Fontijn, Patrick Valentijn, and Harry Fokkens, Archaeology from the Dutch Twilight Zone, in Sophie Bergerbrant and Serena Sabatini (ed), Counterpoint: Essays in Archaeology and Heritage Studies in Honour of Professor Kristian Kristiansen, Oxford: Archaeopress, 2013, 533 , https://www.academia.edu/6072124/Archaeology_of_the_Dutch_Twilight_Zone

[5] David Fontijn, Patrick Valentijn, and Harry Fokkens, Archaeology from the Dutch Twilight Zone, Page 533

[6] Fokkens, Harry, The Periodization of the Dutch Bronze Age: A Critical Review, in Metz, W.H. Beek, B.L. van Steegstra, H (eds) Patina. Essays presented to Jay Jordan Butler on the occasion of his 80th birthday (pp.241-262), Metz, Van Beek & Steegstra
https://www.researchgate.net/publication/28646850_The_periodisation_of_the_Dutch_Bronze_Age_a_critical_review

[7] David Fontijn, Patrick Valentijn, and Harry Fokkens, Archaeology from the Dutch Twilight Zone, Pages 533-534

[8] David Fontijn, Patrick Valentijn, and Harry Fokkens, Archaeology from the Dutch Twilight Zone, Page 535

[9] David Fontijn, Patrick Valentijn, and Harry Fokkens, Archaeology from the Dutch Twilight Zone, Page 535

See also

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CNRS. “Social change may explain decline in genetic diversity of the Y chromosome at the end of the Neolithic period.” ScienceDaily. ScienceDaily, 26 April 2024. www.sciencedaily.com/releases/2024/04/240424111621.htm 

Zeng TC, Aw AJ, Feldman MW. Cultural hitchhiking and competition between patrilineal kin groups explain the post-Neolithic Y-chromosome bottleneck. Nat Commun. 2018 May 25;9(1):2077. doi: 10.1038/s41467-018-04375-6. PMID: 29802241; PMCID: PMC5970157 https://pmc.ncbi.nlm.nih.gov/articles/PMC5970157

[24] mvea, About 7,000 years ago, something weird happened to men: the genetic diversity of their Y chromosomes collapsed. It was as if there was only one man left to mate for every 17 women. The collapse may have been the result of generations of war between patrilineal clans structured around male ancestry, Reddit, https://www.reddit.com/r/science/comments/8nrxzy/about_7000_years_ago_something_weird_happened_to/

Zeng TC, Aw AJ, Feldman MW. Cultural hitchhiking and competition between patrilineal kin groups explain the post-Neolithic Y-chromosome bottleneck. Nat Commun. 2018 May 25;9(1):2077. doi: 10.1038/s41467-018-04375-6. PMID: 29802241; PMCID: PMC5970157. https://pmc.ncbi.nlm.nih.gov/articles/PMC5970157/

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[32] The quote is from:

Sjögren, Karl-Goran, Iñigo Olalde, Sophie Carver, Morten E. Allentoft, Tim Knowles, Guus Kroonen, Alistair W. G. Pike, Peter Schröter, Keri A. Brown, Kate Robson Brown, Richard J. Harrison, Francois Bertemes, David Reich, Kristian Kristiansen, Volker Heyd, Kinship and social organization in Copper Age Europe. A cross-disciplinary analysis of archaeology, DNA, isotopes, and anthropology from two Bell Beaker cemeteries, PLoS ONE 15(11): e0241278, 2020, doi.org/10.1371/journal.pone.0241278

See also:

Kristiansen K et al. , Re-theorising mobility and the formation of culture and language among the Corded Ware Culture in Europe. Antiquity 2017; 91: 334–347 https://www.cambridge.org/core/journals/antiquity/article/retheorising-mobility-and-the-formation-of-culture-and-language-among-the-corded-ware-culture-in-europe/E35E6057F48118AFAC191BDFBB1EB30E

Sjögren K-G, Price TD, Kristiansen K, Diet and mobility in the Corded Ware of central Europe. PLoS One 2016; 11:e0155083 10.1371/journal.pone.0155083  (PubMed) https://pubmed.ncbi.nlm.nih.gov/27223117/

Goldberg A, Günther T, Rosenberg NA, Jakobsson M, Ancient X chromosomes reveal contrasting sex bias in Neolithic and Bronze Age Eurasian migrations. Proc Natl Acad Sci USA 2017; 114/10: 2657–2662. 10.1073/pnas.1616392114 (PubMed) https://pubmed.ncbi.nlm.nih.gov/28223527/

Zeng TC, Aw AJ, Feldman MW, Cultural hitchhiking and competition between patrilineal kin groups explain the post-Neolithic Y-chromosome bottleneck. Nature Comm. 2018; 9: 2077 10.1038/s41467-018-04375-6  (PubMed) https://pubmed.ncbi.nlm.nih.gov/29802241/

Sjögren K-G, Price TD, Kristiansen K, Diet and mobility in the Corded Ware of central Europe. PLoS One 2016; 11:e0155083 10.1371/journal.pone.0155083 
(PubMed) https://pubmed.ncbi.nlm.nih.gov/27223117/

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Goldberg A, Günther T, Rosenberg NA, Jakobsson M, Reply to Lazaridis and Reich: Robust model-based inference of male-biased admixture during Bronze Age migration from the Pontic-Caspian Steppe. Proc Natl Acad Sci USA 2017; 114/20: E3875– E3877. 10.1073/pnas.1704442114 (PubMed) https://pubmed.ncbi.nlm.nih.gov/28476765/

Haak W et al. , Ancient DNA, Strontium isotopes, and osteological analyses shed light on social and kinship organization of the Later Stone Age. Proc Natl Acad Sci USA 2008; 105: 18226–18231. 10.1073/pnas.0807592105 (PubMed) https://pubmed.ncbi.nlm.nih.gov/19015520/

Knipper C et al. , Female exogamy and gene pool diversification at the transition from the Final Neolithic to the Early Bronze Age in central Europe. Proc Natl Acad Sci USA 2017; 114: 10083–10088. 10.1073/pnas.1706355114 (PubMed0)https://pubmed.ncbi.nlm.nih.gov/28874531/

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[34] “The Globulara Amphora culture could possibly have been in contact with the Yamnaya, but that the interaction would mostly have been at the cultural level with very limited migration.“

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See also:

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Francesca Tassi, Stefania Vai, Silvia Ghirotto, Martina Lari, Alessandra Modi, Elena Pilli, Andrea Brunelli, Roberta Rosa Susca, Alicja Budnik, Damian Labuda, Federica Alberti, Carles Lalueza-Fox, David Reich, David Caramelli and Guido Barbujani, Genome diversity in the Neolithic Globular Amphorae culture and the spread of Indo-European languages, 22 November 2017, Proceedings of the Royal Society Biological Sciences, .https://doi.org/10.1098/rspb.2017.1540

Mikhail M. Charniauski et al. (eds.), Eastern exodus of the globular amphora people: 2950-2350 BC. Poznań, Adam Mickiewicz University, Institute of Prehistory 1996, Baltic-Pontic studies 4.

Manfred Woidich, The Western Globular Amphora Culture. A New Model for its Emergence and Expansion, eTopoi Journal of Ancient Sdtudies, , Vol 3, 2014, 67 -85, https://www.topoi.org/wp-content/uploads/2014/08/182-704-1-SM.pdf

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Virginia García‐Díaz, The Domestic Shpere of the Corde Ware Culture, PhD Thesis, Leiden University, 1981, https://collectie.huisvanhilde.nl/pdf/The_domestic_sphere_of_the_corded_ware_culture.pdf#:~:text=The%20Corded%20Ware%20Culture%20(CWC)%20was%20a,Funnelbeaker%20(Trechterbekercultuur%20or%20TRB%20in%20Dutch)%20groups.

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Großmann, Ralph, Interrelations between Corded Ware and Bell Beaker Phenomena?Material Cultures and Identities in the 3rd Millennium BC, in Martin Furholt, Ralph Großmann, Marzena Szmyt, Transitional Landscapes? The 3rd Millennium BC in Europe, Gedruckt mit unterstützung der deutschen forschungsgemeinschaft (DfG), 2016,  Pages 129 – 143, https://www.academia.edu/38915210/R_Großmann_Interrelations_between_corded_ware_and_bell_beaker_phenomena_Material_cultures_and_identities_in_the_3rd_millennium_BC

Sjögren Karl-Göran, Douglas Price T, Kristiansen Kristiansen. Diet and Mobility in the Corded Ware of Central Europe. PLoS One. 2016; 33. https://doi.org/10.1371/journal.pone.0155083 PMID: 27223117

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See also:

Kristiansen, Kristian, The Formation of Language and Culture in Temperate Europe after the Steppe Migrations 3000–2800 BC, Särtryck ur: Årsbok 2017 KVHAA Stockholm 2016 (isbn 978-91-7402-456-2, issn 0083-6796) https://www.vitterhetsakademien.se/download/18.542301d218418c937a71a612/1669133849334/Kristiansen_ÅB_17.pdf

Kristiansen, Kristian, The Formation of Language and Culture in Temperate Europe after the Steppe Migrations 3000–2800 BC, Särtryck ur: Årsbok 2017 KVHAA Stockholm 2016 (isbn 978-91-7402-456-2, issn 0083-6796) https://www.vitterhetsakademien.se/download/18.542301d218418c937a71a612/1669133849334/Kristiansen_ÅB_17.pdf

Furholt, M. Mobility and Social Change: Understanding the European Neolithic Period after the Archaeogenetic Revolution. J Archaeol Res 29, 481–535 (2021). https://doi.org/10.1007/s10814-020-09153-x

“Corded Ware from East to West .” Ancient Europe, 8000 B.C. to A.D. 1000: Encyclopedia of the Barbarian World. . Encyclopedia.com. (July 18, 2025). https://www.encyclopedia.com/humanities/encyclopedias-almanacs-transcripts-and-maps/corded-ware-east-west

[37] Kristiansen, Kristian & Allentoft, Morten & Frei, Karin & Iversen, Rune & Johannsen, Niels & Kroonen, Guus & Pospieszny, Lukasz & Price, T. & Rasmussen, Simon & Sjögren, Karl-Göran & Sikora, Martin & Willerslev, Eske. (2017). Re-theorising mobility and the formation of culture and language among the Corded Ware Culture in Europe. Antiquity. 91. 334-347. 10.15184/aqy.2017.17. https://www.researchgate.net/publication/315990713_Re-theorising_mobility_and_the_formation_of_culture_and_language_among_the_Corded_Ware_Culture_in_Europe

[38] Kristiansen, Kristian, The Formation of Language and Culture in Temperate Europe after the Steppe Migrations 3000–2800 BC, Särtryck ur: Årsbok 2017 KVHAA Stockholm 2016 (isbn 978-91-7402-456-2, issn 0083-6796), Page 124 , htpps://www.vitterhetsakademien.se/download/18.542301d218418c937a71a612/1669133849334/Kristiansen_ÅB_17.pdf

[39] Bourgeois, Quentin & Kroon, Erik, The impact of male burials on the construction of Corded Ware identity: Reconstructing networks of information in the 3rd millennium BC, 12 October 2017, PLOS ONE, 12, 10.1371/journal.pone.0185971, https://www.researchgate.net/publication/320362274_The_impact_of_male_burials_on_the_construction_of_Corded_Ware_identity_Reconstructing_networks_of_information_in_the_3rd_millennium_BC

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[48] Sjögren, Karl-Goran, Iñigo Olalde, Sophie Carver, Morten E. Allentoft, Tim Knowles, Guus Kroonen, Alistair W. G. Pike, Peter Schröter, Keri A. Brown, Kate Robson Brown, Richard J. Harrison, Francois Bertemes, David Reich, Kristian Kristiansen, Volker Heyd, Kinship and social organization in Copper Age Europe. A cross-disciplinary analysis of archaeology, DNA, isotopes, and anthropology from two Bell Beaker cemeteries, PLoS ONE 15(11): e0241278, 2020, doi.org/10.1371/journal.pone.0241278 (pubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC7668604/

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[51] The cemetery of Alburg is located in the city of Straubing, Lower Bavaria, Germany. The cemetery of Irlbach is located in the County of Straubing-Bogen also in Lower Bavaria, Germany. The cemetaries are close to each other on or near the Danube River. If we arbitrarily chose Liège, Belgium as an anchor point for where Griff(is)(es)(ith) YDNA descendants may have lived in the initial stages of this phylogenetic gap, the distance from the cemetaries is approximately 670 kilometers or about 420 miles .

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[54] Zedda Nicolettta, Meheux K, Blöcher J, Diekmann Y, Gorelik AV, Kalle M, Klein K, Titze AL, Winkelbach L, Naish E, Brou L, Valotteau F, Le Brun-Ricalens F, Burger J, Brami M. Biological and substitute parents in Beaker period adult-child graves. Sci Rep. 2023 Oct 31;13(1):18765. doi: 10.1038/s41598-023-45612-3. PMID: 37907573; PMCID: PMC10618162. (PubMed) https://pubmed.ncbi.nlm.nih.gov/37907573/

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Press Release , The Grave’s Embrace: New research sheds light on Bronze Age family relationships, 29 Jan 2024, Johannes Gutenberg Universität Mainz , https://press.uni-mainz.de/the-graves-embrace-new-research-sheds-light-on-bronze-age-family-relationships/

[55] Zedda N, et al. Biological and substitute parents in Beaker period adult-child graves. Sci Rep. 2023 Oct 31;13(1):18765. doi: 10.1038/s41598-023-45612-3. PMID: 37907573; PMCID: PMC10618162. (PubMed) https://pubmed.ncbi.nlm.nih.gov/37907573/

[56] Zedda N, et al. Biological and substitute parents in Beaker period adult-child graves. Sci Rep. 2023 Oct 31;13(1):18765. doi: 10.1038/s41598-023-45612-3. PMID: 37907573; PMCID: PMC10618162. (PubMed) https://pubmed.ncbi.nlm.nih.gov/37907573/

[57] Zedda N, et al. Biological and substitute parents in Beaker period adult-child graves. Sci Rep. 2023 Oct 31;13(1):18765. doi: 10.1038/s41598-023-45612-3. PMID: 37907573; PMCID: PMC10618162. (PubMed) https://pubmed.ncbi.nlm.nih.gov/37907573/

[58] Zedda N, et al. Biological and substitute parents in Beaker period adult-child graves. Sci Rep. 2023 Oct 31;13(1):18765. doi: 10.1038/s41598-023-45612-3. PMID: 37907573; PMCID: PMC10618162. (PubMed) https://pubmed.ncbi.nlm.nih.gov/37907573/

[59] Olalde I, Brace S, Allentoft ME, et al., The Beaker phenomenon and the genomic transformation of northwest Europe. Nature. 2018 Mar 8;555(7695):190-196. doi: 10.1038/nature25738. Epub 2018 Feb 21. Erratum in: Nature. 2018 Mar 21;555(7697):543. doi: 10.1038/nature26164. PMID: 29466337; PMCID: PMC5973796. (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC5973796/

[60] Grossmann, Ralph. “R. Großmann, Interrelations between Corded Ware and Bell Beaker Phenomena? Material Cultures and Identities in the 3rd Millennium BC.” Furholt u. a. 2016 M. Furholt/R. Großmann/M. Szmyt (Hrsg.), Transitional Landscapes? The 3rd Millennium BC in Europe: Proceedings of the International Workshop “Socio-Environmental Dynamics over the Last 12000 Years: the Creation of Landscapes III (15th-18th April 2013)” in Kiel., vol. 9, 2016. https://www.academia.edu/38915210/R_Großmann_Interrelations_between_corded_ware_and_bell_beaker_phenomena_Material_cultures_and_identities_in_the_3rd_millennium_BC

Beckerman, Sandra. “Dutch Beaker Chronology Re-Examined. .” Palaeohistoria 2011/2012, Vol 53-54., 2011/2012, Groningen: University of Groningen. 25-64. https://www.academia.edu/2304446/Dutch_beaker_chronology_re_examined

Harry Fokkens, Background to Dutch Beakers: A Critical Review of the Dutch Model, in Harry Fokkens & Freanco Nicolis, eds, Background to Beakers Inquiries into regional cultural backgrounds of the Bell Beaker Complex, Sidestone, 2012 https://www.academia.edu/5507545/Background_to_Dutch_Beakers

[61] Fokkens, Harry, Background to Dutch Beakers. A critical review of the Dutch model, in Harry Fokkens & Franco Nicolis, eds, Background to Beakers Inquiries in regional cultural backgrounds of the Bell Beaker Complex, Leiden: Sidestone Press, 2012, 9-36, https://woolmerforest.org.uk/E-Library/B/Background%20to%20Beakers%20-%20Inquiries%20in%20regional%20cultural%20backgrounds%20of%20the%20Bell%20Beaker%20Complex.pdf

[62] Fokkens, Harry, Background to Dutch Beakers. A critical review of the Dutch model, in Harry Fokkens & Franco Nicolis, eds, Background to Beakers Inquiries in regional cultural backgrounds of the Bell Beaker Complex

Davidski, Dutch Beakers: like no other Beakers, 19 Jan 2019, EuroGenes Blog, https://eurogenes.blogspot.com/2019/01/dutch-beakers-like-no-other-beakers.html

[63] Fokkens, Harry, Background to Dutch Beakers. A critical review of the Dutch model, in Harry Fokkens & Franco Nicolis, eds, Background to Beakers Inquiries in regional cultural backgrounds of the Bell Beaker Complex

Davidski, Dutch Beakers: like no other Beakers, 19 Jan 2019, EuroGenes Blog, https://eurogenes.blogspot.com/2019/01/dutch-beakers-like-no-other-beakers.html

[64] Olalde I, et al., The Beaker phenomenon and the genomic transformation of northwest Europe. Nature. 2018 Mar 8;555(7695):190-196. doi: 10.1038/nature25738. Epub 2018 Feb 21. Erratum in: Nature. 2018 Mar 21;555(7697):543. doi: 10.1038/nature26164. PMID: 29466337; PMCID: PMC5973796. https://pmc.ncbi.nlm.nih.gov/articles/PMC5973796/

[64a] Vander Linden M. For Whom the Bell Tolls: Social Hierarchy vs Social Integration in the Bell Beaker Culture of Southern France (Third Millennium bc). Cambridge Archaeological Journal. 2006; 16(3):317-332. doi: 10.1017/S0959774306000199, https://www.cambridge.org/core/journals/cambridge-archaeological-journal/article/abs/for-whom-the-bell-tolls-social-hierarchy-vs-social-integration-in-the-bell-beaker-culture-of-southern-france-third-millennium-bc/CB0BA59778DC57F42C4ECD5FD667786C ; also found at: https://www.academia.edu/840057/For_Whom_the_Bell_Tolls_Social_Hierarchy_vs_Social_Integration_in_the_Bell_Beaker_Culture_of_Southern_France_Third_Millennium_small_BC_small_

Linden MV. What linked the Bell Beakers in third millennium BC Europe? Antiquity. 2007; 81(312): 343-352. doi: 10.1017/S0003598X00095223, https://www.cambridge.org/core/journals/antiquity/article/abs/what-linked-the-bell-beakers-in-third-millennium-bc-europe/52D01B1D5AA2631C4D4FB06A6CF151E7

Racimo, F., Sikora, M., Vander Linden, M., Schroeder, H. and Lalueza-Fox, C., 2020. Beyond broad strokes: sociocultural insights from the study of ancient genomes. Nature Reviews Genetics, 21, 355-366 , https://eprints.bournemouth.ac.uk/33588/1/Review%20ancient%20DNA%20-%20Fernando%2C%20Martin%2C%20Marc%2C%20Hannes%2C%20Carles.pdf

[65] David Fontijn, Patrick Valentijn, and Harry Fokkens, Archaeology from the Dutch Twilight Zone, in Sophie Bergerbrant and Serena Sabatini (ed), Counterpoint: Essays in Archaeology and Heritage Studies in Honour of Professor Kristian Kristiansen, Oxford: Archaeopress, 2013, 531-540 , https://www.academia.edu/6072124/Archaeology_of_the_Dutch_Twilight_Zone

Großmann, Ralph, Interrelations between Corded Ware and Bell Beaker Phenomena?Material Cultures and Identities in the 3rd Millennium BC, in Martin Furholt, Ralph Großmann, Marzena Szmyt, Transitional Landscapes? The 3rd Millennium BC in Europe, Gedruckt mit unterstützung der deutschen forschungsgemeinschaft (DfG), 2016,  Pages 129 – 143, https://www.academia.edu/38915210/R_Großmann_Interrelations_between_corded_ware_and_bell_beaker_phenomena_Material_cultures_and_identities_in_the_3rd_millennium_BC

Bell Beaker Culture, Wikipedia, This page was last edited on 21 June 2025, https://en.wikipedia.org/wiki/Bell_Beaker_culture

[66] “Bell Beaker people took advantage of transport by sea and rivers, creating a cultural spread extending from Ireland to the Carpathian Basin and south along the Atlantic coast and along the Rhône valley to Portugal, North Africa, and Sicily, even penetrating northern and central Italy“

Bell Beaker Culture, Wikipedia, This page was last edited on 11 August 2025, https://en.wikipedia.org/wiki/Bell_Beaker_culture

Van de Noort, Robert, Exploring Agency Behind the Beaker Phenomenon The navigator’s tale, in Harry Fokkens & Franco Nicolis, eds, Inquiries in Regional Cultural Backgrounds of the Bell Beaker Complex, leden: Sidestone Press, 2021, 61-80 , https://www.sidestone.com/openaccess/9789088900846.pdf

[67] “ People of the Bell Beaker tradition are characterized as being sedentary agriculturalists with supplementary animal husbandry, living in small villages of probably not more than thirty to fifty inhabitants, with a form of social organization based on classes. Settlements were located on good agricultural soils, near a source of water such as a river estuary, commanding a valley route, or sited in a strong defensive position, such as the walled and fortified communities of Zambujal and Vila Nova de São Pedro. These fortified settlements, of which Zambujal is a good example, may have functioned as production and trade centers, manufacturing raw products into finished goods which were then traded. The fortifications were therefore necessary to protect the area from raiding activities of other groups. “

“Bell Beaker houses tend to be circular or oval in shape, up to 7 m. in diameter, and sometimes accompanied by adjacent structures which may have served as animal pens. Rectangular houses of relatively large size (6 by 20 m.) have also been noted for Bell Beaker areas in the Netherlands and in Great Britain. In these closely set posts served to support roof and walls. Generally round or oval houses were made of adobe bricks or flat stones, as at Zambujal, and were probably occupied by a single nuclear family. Differences in wealth and status seem to be indicated by the size and complexity of the individual dwellings.“

Beierle, John, Traditional Summary: Bell Beaker, Bell Beaker E050, eHRAF Archaeology, https://ehrafarchaeology.yale.edu/traditions/e050/summary

See also:

Czebreszuk, Janusz, “Bell Beakers from West to East .” Ancient Europe, 8000 B.C. to A.D. 1000: Encyclopedia of the Barbarian World. . Encyclopedia.com. 14 Aug. 2025 https://www.encyclopedia.com/humanities/encyclopedias-almanacs-transcripts-and-maps/bell-beakers-west-east#:~:text=This%20area%20has%20yielded%20many,combined%20into%20a%20unique%20whole.

Bell Beaker Culture, Wikipedia, This page was last edited on 29 August 2025, https://en.wikipedia.org/wiki/Bell_Beaker_culture

David Reich Lab, Supplementary Information, doi:10.1038/nature25738, https://reich.hms.harvard.edu/sites/reich.hms.harvard.edu/files/inline-files/41586_2018_BFnature25738_MOESM2_ESM.pdf

[68] See the various citations that are referenced in prior sections of this story.

[69] ” On the basis of the analysis of settlement patterns, house types, and mortuary goods, Bell Beaker social organization seems to have been distinguished by two major features: (1) occupational status determined by craft specialization, and (2) the division of society into classes of non-elites and elites, the latter based on wealth and the ability to control the distribution of high-status objects in the society. With the spread of the beaker tradition into western Europe, accompanied by the widespread use of copper and gold in societies where they were previously unknown, the development of new technologies and the increased need for novel luxury items formed the basis for the eventual establishment of chiefdoms with their characteristically ranked or hierarchical societies. ”

” A form of loose political control in Bell Beaker society was maintained by the wealthy elite, who, could summon on demand both a labor and a protective military force as needed. This form of political power approached that which we would call a chiefdom, or at least an organization very similar to it. “

Beierle, John, Traditional Summary: Bell Beaker, Bell Beaker E050, eHRAF Archaeology, https://ehrafarchaeology.yale.edu/traditions/e050/summary

[70] “… (S)ingle burial, communal burial, and reuse of Neolithic burial sites are found throughout the Bell Beaker zone. This overturns a previous conviction that single burial was unknown in the early or southern Bell Beaker zone, and so must have been adopted from Corded Ware in the contact zone of the Lower Rhine, and transmitted westwards along the exchange networks from the Rhine to the Loire,  and northwards across the English Channel to Britain.”

 Bell Beaker Culture, Wikipedia, This page was last edited on 11 August 2025, https://en.wikipedia.org/wiki/Bell_Beaker_culture

See also:

Zedda, N., Meheux, K., Blöcher, J. et al. Biological and substitute parents in Beaker period adult–child graves. Sci Rep 13, 18765 (2023). https://doi.org/10.1038/s41598-023-45612-3

Sjögren, Karl-Goran, et al, Kinship and social organization in Copper Age Europe. A cross-disciplinary analysis of archaeology, DNA, isotopes, and anthropology from two Bell Beaker cemeteries, PLoS ONE 15(11): e0241278, 2020, doi.org/10.1371/journal.pone.0241278 (pubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC7668604/

Torben Sarauw, Male symbols or warrior identities? The ‘archery burials’ of the Danish Bell Beaker Culture, Journal of Anthropological Archaeology, Volume 26, Issue 1, 2007, Pages 65-87, ISSN 0278-4165, https://doi.org/10.1016/j.jaa.2006.05.001.
https://www.sciencedirect.com/science/article/pii/S0278416506000365

[71] ” Despite the fact that many thousands of graves have been excavated, it is not clear that there were any institutionalized high status groups in Bell Beaker Europe. The reason for this is that the complex as it is known from graves is characterized by a striking uniformity in the set of grave goods. This does not exclude the existence of apparently important individuals …  or other seemingly high status individuals buried with gold objects in Central Europe and Iberia. It is also important to recognize the possibility of an ascertainment bias: the Beaker burials that are best characterized, some of which discovered underneath burial mounds, may represent an exclusive segment of society. Some archaeologists have suggested that the burials of the Beaker Complex represent a social elite that introduced new customs and technologies. It is plausible that the majority were buried without elements of the Beaker package and therefore are less recognizable in archaeological context. ”

David Reich Lab, Supplementary Information, doi:10.1038/nature25738, https://reich.hms.harvard.edu/sites/reich.hms.harvard.edu/files/inline-files/41586_2018_BFnature25738_MOESM2_ESM.pdf

See also:

Beierle, John, Traditional Summary: Bell Beaker, Bell Beaker E050, eHRAF Archaeology, https://ehrafarchaeology.yale.edu/traditions/e050/summary

Clément, Nicolas, The prestige of warriors: Bell Beaker archers’ equipment in Central Europe, Préhistoires Méditerranéennes [En ligne], 8 | 2020, mis en ligne le 29 janvier 2021, consulté le 28 août 2025. URL : http://journals.openedition.org/pm/2167  ; DOI : https://doi.org/10.4000/pm.2167

[72] Lemercier, Olivier (December 2018). “Think and Act: Local Data and Global Perspectives in Bell Beaker Archaeology”. Journal of Neolithic Archaeology. 20 (Special Issue 4): 77–96. doi:10.12766/jna.2018S.5 , https://www.academia.edu/38019997/LEMERCIER_O_2018_Think_and_Act_Local_Data_and_Global_Perspectives_in_Bell_Beaker_Archaeology_Journal_of_Neolithic_Archaeology_20_2018S_p_77_96

Lemercier, Olivier, Historical model of settling and spread of Bell Beakers Culture in the mediterranean France, Symposium of the Northeast Frontier of Bell Beakers – Poznan (Poland) – 26-29 May 2002. Session : What’s new in the study of Bell Beakers after Riva del Garda. https://shs.hal.science/halshs-00087311/file/Historical_Model_Bell_Beakers_-_2004.pdf

Vander Linden, Marc, ‘Bell Beaker Pottery and Society’, in Chris Fowler, Jan Harding, and Daniela Hofmann (eds), The Oxford Handbook of Neolithic Europe (2015; online edn, Oxford Academic, 5 Dec. 2014), https://doi.org/10.1093/oxfordhb/9780199545841.013.031

Favrel, Quinten, Around Maritime beaker: the vases à cordon, linear beakers and épicampaniforme productions in north‑western France, 16 Jan 2020, OpenEdition Journals, https://journals.openedition.org/pm/2163?lang=en

Drenth, Erik and Mom, Vincent, Continuity and change: on a computer-aided classification of Late Neolithic and Early Bronze Age pottery from the Netherlands, 1. Jan 1955, Computer Applications to Archaeology 2009 Williamsburg, Virginia, USA. March 22-26, 2009, https://www.researchgate.net/publication/242457506_Continuity_and_change_on_a_computer-aided_classification_of_Late_Neolithic_and_Early_Bronze_Age_pottery_from_the_Netherlands

J. D. Van Der Waals, J.D., W. Glasbergen, Beaker Types and their Distribution in the Netherlands, Paleohistoria, 4 , 1955, 1- 630https://ugp.rug.nl/Palaeohistoria/article/view/24772/22220

[73] Dalal, Vasundhra; Pasupuleti, Nagarjuna; Chaubey, Gyaneshwer; Rai, Niraj; Shinde, Vasant Advancements and Challenges in Ancient DNA Research: Bridging the Global North–South Divide. Genes 2023, 14, 479. https://doi.org/10.3390/genes14020479

Briggs, Lisa, Ancient DNA Research in Maritime and Underwater Archaeology: Pitfalls, Promise, and Future Directions, in Maritime and Underwater Archaeology: Pitfalls, Promise, and Future Directions’, Open Quaternary, 6(1), p. 3. https://openquaternary.com/articles/10.5334/oq.71

Aja Golob, Pia Kravanja, Monica Concato, Tamara Leskovar, Irena Zupanič Pajnič,
Searching for alternative high DNA-yielding bone types for DNA analysis of aged skeletal remains, Forensic Science International, Volume 362, 2024, 112184,
ISSN 0379-0738, https://doi.org/10.1016/j.forsciint.2024.112184 .
(https://www.sciencedirect.com/science/article/pii/S0379073824002652 )

Latham Krista E, Miller Jessica J. DNA recovery and analysis from skeletal material in modern forensic contexts. Forensic Sci Res. 2018 Oct 8;4(1):51-59. doi: 10.1080/20961790.2018.1515594. PMID: 30915417; PMCID: PMC6427720 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC6427720/

Gaeta Raffaele. Ancient DNA and paleogenetics: risks and potentiality. Pathologica. 2021 Apr;113(2):141-146. doi: 10.32074/1591-951X-146. PMID: 34042097; PMCID: PMC8167392 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC8167392/

Iñigo Olalde, Eveline Altena, Quentin Bourgeois, Harry Fokkens, Luc Amkreutz, Marie-France Deguilloux, Alessandro Fichera, Damien Flas, Francesca Gandini, Jan F. Kegler, Lisette M. Kootker, Kirsten Leijnse, Leendert Louwe Kooijmans, Roel Lauwerier, Rebecca Miller, Helle Molthof, Pierre Noiret, Daan C. M. Raemaekers, Maïté Rivollat, Liesbeth Smits, John R. Stewart, Theo ten Anscher, Michel Toussaint, Kim Callan, Olivia Cheronet, Trudi Frost, Lora Iliev, Matthew Mah, Adam Micco, Jonas Oppenheimer, Iris Patterson, Lijun Qiu, Gregory Soos, J. Noah Workman, Ceiridwen J. Edwards, Iosif Lazaridis, Swapan Mallick, Nick Patterson, Nadin Rohland, Martin B. Richards, Ron Pinhasi, Wolfgang Haak, Maria Pala, David Reich, Long-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups, bioRxiv 2025. 03.24.644985; doi: https://doi.org/10.1101/2025.03.24.644985

Fokkens, H., Steffens, B. J. W. & van As, S. F. . Farmers, fishers, fowlers, hunters: knowledge generated by development-led archaeology about the Late Neolithic, the Early Bronze Age and the start of the Middle Bronze Age (2850–1500 cal BC) in the Netherlands. Ned. Archeol. Rapp. 53, 978–990 (2016) https://www.academia.edu/31484011/Farmers_fishers_fowlers_hunters_Knowledge_generated_by_development_led_archaeology_about_the_Late_Neolithic_the_Early_Bronze_Age_and_the_start_of_the_Middle_Bronze_Age_2850_1500_cal_BC_in_the_Netherlands

[74] Iñigo Olalde,et alLong-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups, bioRxiv 2025. 03.24.644985; doi: https://doi.org/10.1101/2025.03.24.644985

Ibrahim, Jamal, Vlad Brumfeld, Yoeseph Addadi, Sarah rubin, Steve Weiner, Elisabetta Boaretto, The petrous bone contains high concentrations of osteocytes: One possible reason why ancient DNA is better preserved in this bone, Published: October 25, 2022, https://doi.org/10.1371/journal.pone.0269348

Hansen HB, Damgaard PB, Margaryan A, Stenderup J, Lynnerup N, Willerslev E, et al. (2017) Comparing Ancient DNA Preservation in Petrous Bone and Tooth Cementum. PLoS ONE 12(1): e0170940. https://doi.org/10.1371/journal.pone.0170940

Hansen HB, Damgaard PB, Margaryan A, Stenderup J, Lynnerup N, Willerslev E, Allentoft ME. Comparing Ancient DNA Preservation in Petrous Bone and Tooth Cementum. PLoS One. 2017 Jan 27;12(1):e0170940. doi: 10.1371/journal.pone.0170940. PMID: 28129388; PMCID: PMC5271384 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC5271384/

Renaud G, Schubert M, Sawyer S, Orlando L. Authentication and Assessment of Contamination in Ancient DNA. Methods Mol Biol. 2019;1963:163-194. doi: 10.1007/978-1-4939-9176-1_17. PMID: 30875054 (PubMed) https://pubmed.ncbi.nlm.nih.gov/30875054/

[75] Iñigo Olalde,et alLong-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups, bioRxiv 2025. 03.24.644985; doi: https://doi.org/10.1101/2025.03.24.644985

[76] Iñigo Olalde,et alLong-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups, bioRxiv 2025. 03.24.644985; doi: https://doi.org/10.1101/2025.03.24.644985

[77] Iñigo Olalde,et alLong-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups, bioRxiv 2025. 03.24.644985; doi: https://doi.org/10.1101/2025.03.24.644985

[78] Iñigo Olalde,et alLong-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups, bioRxiv 2025. 03.24.644985; doi: https://doi.org/10.1101/2025.03.24.644985

This story is the second of a four part story focusing on the second major gap in haplogroup subclades in the G Haplogroup phylogenetic tree of the Griff(is)(es)(ith) patrilineal line of descent. The first part of this story focused on the unique nature of the Meuse – Rhine River watershed and possible environmental influences on the lack of subclades in this phylogenetic gap.

This second part of the story focuses on:

• the three haplogroups between the two major phylogenetic gaps;
• the unique nature of the Mesuse and Rhine River watershed on haplogroup migration; and
• examining possible social-cultural influences in the neolithic age that may explain the lack of identified subclades (ancestors) in the migratory path of the Griffis genetic paternal line during this time period.

The 2,850 year Gap between G-FGC7516 and G-Z6748: This phylogenetic gap was associated with haplogroup G-FGC7516. This common ancestor was born around 2200 BCE. The next genetic ancestor on the Griff(is)(es)(ith) YDNA line was associated with the genetic SNP mutation defining the G-Z6748 haplogroup, 2,850 years later. This gap represents about 95 generations.

There are a number of social, cultural and demographic factors that may have contributed to conditions that either limited the proliferation of G haplogroup subclades or were unfavorable for the preservation or detection of intermediate lineages over nearly three millennia. The convergence of bronze age male bottlenecks, social kinship structures that hindered G haplogroup proliferation, possible border conflicts between social groups along the Rhine River, late-Roman demographic collapse, female-biased assimilation practices, environmental instability in the delta landscape, and ‘just simply basic’ research sampling gaps can collectively explain the missing subclades.

The Migratory Path of Genetic Ancestors and Socio-Cultural Associations

The ancestors that are associated with the beginning of this phylogenetic gap may have lived when the Michelsberg culture, Funnelbeaker culture, the Corded Ware or Bell Beaker cultures existed. The ancestor associated with the G-FGC477 haplogroup may have lived when local bronze-Iron age tribes in the Meuse-Rhine watershed were predominate (see table one).

Haplogroups that precede the phylogenetic gap discussed in this part of the story are identified in italicized purple in table one.

Table One: Most Recent Common Ancestors (MCRA) and Possible Cultural Associations

Haplogroup	MRCA Birth Date	MRCA Birth Date Range 68 % Confidence Interval	Possible Cultural Associations Based on Birth Range & Geographic Location
G-Z1900	4600 BCE	5129 – 4191 BCE	Linear Pottery, Cortaillod, Pfyn, Lake-Dwelling (Pile Dweller), Stein Group, Funnelbeaker
G-Z6901	3950 BCE	4420 – 3572 BCE	Horgen, Michelsberg, Funnelbeaker
G-Z1817	3100 BCE	3470 – 2743 BCE	Funnelbeaker, Corded Ware
G-Z727	2550 BCE	2882 – 2229 BCE	Funnelbeaker, Corded Ware, Unetice
G-FGC477	2250 BCE	2615 – 1929 BCE	Corded Ware, Bell Beaker, Unetice
G-FGC7516	2200 BCE	2555 – 1876 BCE	Corded Ware, Bell Beaker, Unetice, Hilversum
Missing Haplogroups			Hilversum, Unetice, Tumulus, Urnfield, Hallstatt, La Tène, Mediomatrici
G-Z6748	650 CE	519 – 788 CE	German and Celtic Tirbal cultures: Hallstatt, La Tène, Mediomatrici

The missing genetic generations associated within the phylogenetic gap may have lived during the presence of the Hilversum culture (1850-1650 BCE), Unetice culture (2,300-1,600 BCE), the Tumulus culture (1,600-1,200 BCE), and the Urnfield culture (1,300-750 BCE).

Many of the missing genetic generations within the phylogenetic gap could have been part of Celtic cultures, with the Hallstatt culture (800-450 BCE) giving way to the La Tène culture (450-50 BCE) and the Mediomatrici, from the Hallstatt tradition, moved in the 9th century BCE to settle west of the Rhine, with their territory extending to Speyer in the Rhineland-Palatinate.

The ancestor associated with the haplogroup G-Z6748 at end of the gap could have been associated with Roman or post Roman transitions or German tribal societies or post Roman tribal cultures.

Avoiding Ecological Fallacies

While I discuss various cultural groups that may have existed and coincided with the estimated migratory path of the Griff(is)(es)(ith) genetic paternal line, one should be cautious about making claims about individuals or characteristics of the migratory path based on group-level trends. One needs to always consider the possibility of exceptions and diversity when analyzing individual genealogical data and placing that data within the context of larger historical groups.

Illustration One: Ecological Fallacy

This level of caution avoids the pitfalls of an ecological fallacy. An ecological fallacy is a logical and statistical error that occurs when conclusions about individuals are drawn from group-level data. In genetics, this means assuming that patterns or characteristics observed in large populations or cultural groups necessarily apply to each individual within those groups. [1]

When analyzing individual YDNA migration patterns using broad geographical or historical data associated with ancient cultural groups, the ecological fallacy can arise in several ways:

• Attributing Group Trends to Individuals: Macro-level data (e.g., the prevalence of a YDNA haplogroup in an ancient population) may not reflect the ancestry or migration history of a specific individual. For example, just because a haplogroup is common in a region historically associated with a particular culture does not mean every individual with that haplogroup shares the same cultural or migratory background.
• Overlooking Genetic Diversity: Ancient populations were often genetically diverse, and cultural boundaries rarely matched genetic ones. Inferring that an individual’s YDNA reflects the migration or history of an entire cultural group ignores this diversity and the possibility of admixture, founder effects, or population bottlenecks.
• Ignoring Subgroup Variation: Macro-level data can mask important substructures within populations. For instance, two individuals from the same region and time period may have very different YDNA lineages due to local migrations, social structure, or random genetic drift.

Migration of Known Haplogroups in between the Two Phylogenetic Gaps

It is interesting to discuss information related to the three known genetic paternal ancestors that are documented between the two phylogentic gaps: haplogroups G-CTS9737, G-Z1900 and G-Z6901. The three haplogrops are between haplogroup G-L497, the endpoint of the first phylogenetic gap, and the begining of the second phylogenetic gap which is represented by a pair of haplogroups G-Z7648 and G-Z727 (see illustration two). The haplogroup G-Z6901 is not depicted in illustration two. There is not enough reported geographical information to estimate its ancestral location.

Illustration Two: Haplogroups in between the Two Phylogenetic Gaps

As reflected in column two in table two below, the three haplogroups are relatively close to each other based on the estimated birth of the ancestor associated with each haplogroup. The time elapsed in red italicized type represent the two major phylogenetic gaps

Table Two: Haplogroups in between the Two Phylogenetic Gaps

Haplogroup	Estimated Birth Date (BCE)	Time Elapsed from prior Haplogroup (Years)	Immediate Descendants (No. of Subclades)	Ancient Samples *
G-Z6748	650 (CE)	2,850	2	– –
G-FGC477	2250	300	2	6
G-Z727	2550	550	3	59
G-Z1817	3100	850	2	62
G-6901	3950	700	1	66
G-Z1900	4650	250	2	72
G-CTS9737	4900	600	1	76
G-L497	5500	3,450	2	84

As the Griff(is)(es)(ith) patrilineal ancestors migrated westward from the eastern areas of the Danube river, the influences of the of the Linear Pottery culture (LBK) were more evident. The name LBK derives from an abbreviation of the German Linienbandkeramik. [2] The LBK was prominent along the Danube River from approximately 5500 BC to 4500 BC. The culture originated on the middle Danube, particularly in regions of western Hungary, and spread westward along the Danube valley into Central Europe, including present-day Austria, Slovakia, and Germany. This period marks the initial spread of agriculture in Europe, with the LBK representing a major Neolithic horizon in the region. The earliest phase began around 5500 BC, and the culture persisted in various local forms until about 4500 BC. [3]

The first phylogenetic gap in the migratory path of the Griff(is)(es)(ith) family line mirrored the general migratory path of the Early European Farmers along the Danube river valley. As depicted in illustration three, the generations that were part of the tail end of the first phylogentic gap probably were part of the Western Linear Pottery culture.

(The LBK) “began in regions of densest occupation on the middle Danube (Bohemia, Moravia, Hungary) and spread over about 1,500 km (930 mi) along the rivers in 360 years. The rate of expansion was therefore about 4 km (2.5 mi) per year“. [4]

Illustration Three: European Cultures in the Middle Neolithic ( 5500 —  4500 BCE) and the Migratory Path of the Griff(is)(es)(ith) Paternal Famly Line

As discussed in the first part of this story, there is strong archaeological evidence that the LBK culture practiced patrilocality (males remaining in their birth communities) and patrilineal wealth inheritance. This pattern aligns with a patrilocal kinship system. [5] Genetic analyses of LBK populations show Y-chromosome haplogroups (e.g., G2a and I2) passed through male lines, consistent with patrilineal descent. Conversely, there is archealogical evidence of mitochondrial DNA diversity which suggests female exogamy. Female exogamy refers to the social practice where women marry outside of their own social group or community. [6]

Archaelogical evidence collectively underscores a society where male lineages dictated resource access and social standing, with women integrating into new communities through marriage. The incremental impact over successive generations of these cultural practices may have resulted in a phylogenetic tree for the Griff(is)es)(ith) paternal line characterized by long, narrow branches during this transformative historical period, reflecting limited genetic diversity despite the geographical and population expansion of the G haplogroup into central and western Europe. The implication of this observation is that the male ancestors of the Griff(s)(es)(ith) genetic line were likely not in positions or in kinship groups of power in various historical cultural contexts.

The most recent common ancestor (MRCA) that is at the end of the first phylogenetic gap is associated with the G-L497 haplogroup. It is estimated that the MRCA of G-L497 resided in the Tyrolean Alps. [7] As indicated in illustration four, based on a 68 percent confidence interval, it is likely that the common ancestor associated with the G-L497 haplogroup was born between 6087 BCE and 5028 BCE. It is estimated that he was born around 5522 BCE.

Illustration Four: Scientific Details for Birth for the Most Recent Common Ancestor Associated with Haplogroup G-L497

G-L497 is a significant Y-chromosome haplogroup. It represents one of the major European branches of haplogroup G. [8] The earliest ancient DNA samples bearing G-L497 have been found in Baden, Řivnáč, and Únětice cultures (Central Europe), as well as in Copper Age northern Italy and Bronze Age southeastern France. [9] Haplogroup G-L497 shows a distinct European concentration, particularly along the Rhine River and in alpine regions. The haplogroup reaches its highest frequencies in the Tyrolean Alps, where some valleys show concentrations above 40 percent. [10]

Perhaps when the most recent common ancestor associated with the G-L497 haplogroup was part of early waves or groups of early european farmers that lived in the Tyrolean region. During 6000–5100 BCE this region sat at a crossroads of Mesolithic and Neolithic traditions. The earliest communities were semi-nomadic hunters, gradually replaced or assimilated by incoming farming groups associated with the Impressed Ware/Cardial and Linear Pottery cultures. This transition laid the foundations for permanent settlement and more complex societies in the Alpine area by the late Neolithic period.

Table Three: Phases and Cultures in the Tyrolean Area Around 6000 and 5000 BCE

Period	Main Cultural Group	Key Features	Region
Late Mesolithic	Local Hunter-Gatherers	Mobile, stone tools, wild fauna	Throughout Tyrol
Early Neolithic	Impressed Ware/Cardial	Pottery with shell impressions, early farming	Southern Tyrol /Adjacent Alps
Early Neolithic	Linear Pottery (LBK)	Longhouses, decorated pottery, cereal agriculture	North Alpine fringe

Between 6000 BCE and 5000 BCE, the Tyrolean region (encompassing present-day North and South Tyrol in the Alps) experienced a significant transition from Mesolithic hunter-gatherer lifestyles to early Neolithic farming communities. Evidence from archaeology and genetics reveals two primary cultural phases during this era. [11] Illustration five indicates the approximate area where the common ancestor associated with the G-L497 haplogroup may have lived during this expansion.

Illustration Five: Expansion of Farming in Western Eurasia

As indicated in table two above, the documented haplogroups between the two phylogenetic gaps were relatively close together in terms of chronological time. In addition there are relatively more ancient DNA samples associated with the three haplogroups. The migratory movement of the Griff(is)(es)(ith) genetic paternal line may have slowed down a bit during this time period. The estimated birthdates of the MRCA subclades for CTS9737, Z1900 and Z6901 are 4900 BCE, 4650 BCE and 3950 BCE respectively. The statistical 68 percent confidence intervals for when they were likely born overlap. (see illustrations six through eight).

Illustration Six: Scientific Details for Birth for the Most Recent Common Ancestor Associated with Haplogroup G-CTS9737

Illustration Seven: Scientific Details for Birth for the Most Recent Common Ancestor Associated with Haplogroup G-Z1900

Illustration Eight: Scientific Details for Birth for the Most Recent Common Ancestor Associated with Haplogroup G-Z6901

During the period roughy between 5000 and 3000 BCE, present-day Switzerland was home to a range of Neolithic cultures characterized by settled farming, animal husbandry, and village life. The archaeological evidence reveals several key cultural groups and settlement types that shaped the region’s prehistory. Based on when the haplogroups G-CTS9737 and G-Z1900 lived in the Austrian – Swiss area, they may have lived in areas inhabited by the Linear Pottery, Cortailod, Pfyn, or Lake-Dwelling cultures.

Table Four: Cultural Groups in Neolithic Swtzerland

Possible Griff(is)(es)(ith) Haplogroup	Culture/Group	Date (BCE)	Area	Key Features
G-CTS9737 G-Z1900	Linear Pottery	~ 5500–4500	Swiss Plateau	Early farming, pottery
G-Z1900	Cortaillod	~ 4300–3500	West/Central	Advanced pottery, lake dwellings
G-Z1900	Pfyn	~ 3900–3500	East	Incised pottery, villages
G-Z1900	Lake-Dwelling	~ 5000 onward	Lakeshore regions	Stilt houses, agro-pastoralism
– – –	Horgen	~ 3500-2850	North Switzerland	Simple pottery, robust tools, lakeshore villages
– – –	Schnurkeramik	~ 2700 – 2400	Lowlands	Distinctive battle axes, wodden wheels, use of copper, small grave mounds

The Cortailod culture flourished approximately between 4300 and 3500 BCE, primarily in western and central Switzerland. It was known for lakeside settlements (pile dwellings), distinctive pottery styles, and specialized bone and stone tools. Archaeological finds are notably concentrated around Lake Neuchâtel. [12]

“As archaeological methods and understanding progressed, this single, large culture complex was broken down into four key phases: the Cortaillod, the Pfyn, the Horgen, and the Schnurkeramik. Each was particular to a different region of Switzerland, and while there was some overlap between one culture and the next, each represents a particular time period, region, and culture.” [13]

Table Five: Evidence of Patrilocaliy and Patrilinealty in Neolithic Swiss Cultures

Culture	Evidence for Patrilocality	Evidence for Patrilineality	Strength of Evidence
Linear Pottery	Genetic & isotope studies	Kin groupings & artifacts	Strong
Cortaillod	Not directly documented	Not directly documented	Lacking due to poor mortuary evidence
Pfyn	Genetic & isotope studies	Likely, through genetic continuity	Strong for patrilocality, fairly strong for patrilineality

The Pfyn culture was a contemporary with the Cortaillod culture, mainly in eastern Switzerland, dating to roughly 3900–3500 BCE. It was characterized by pottery with incised and impressed decoration, small-scale settlements, and farming communities. [14]

The lake dwelling pile dweller communities were Neolithic societies that built wooden stilt houses along lakeshores and marshlands. These pile dwellings are especially well-preserved around Lake Zurich, Lake Geneva, and Lake Constance. The archeological evidence asociated with these communities offer deep clues of their technology and social organization. [15]

As previously mentioned, genetic and isotopic studies of individuals from Linear Pottery (LBK) cemeteries across Central Europe—including the western reaches into Switzerland—consistently show that males generally had local isotopic signatures while females exhibited more variation. This pattern indicates that men tended to stay in their birthplace while women married into outside groups from other regions, which is strong evidence for patrilocal residence customs. [16]

Genetic analyses revealed that kinship groups within settlements were centered around paternal lines. In particular, there is significantly less variance in geographic origin among males than females, and often males buried with indicative items (such as axes or adzes) were related, supporting a tendency toward patrilineal descent and inheritance. [17]

Direct evidence for the internal kinship or residence structures (patrilocal or otherwise) for the Cortailod culture is sparse, largely due to the very limited data on funerary practices and the absence of detailed genetic studies. Archaeological literature repeatedly highlights the major gap in our understanding of family and burial customs for the Cortaillod, making it difficult to confirm their social organization. While there is robust evidence for patrilocality from both earlier (LBK) and later (Pfyn and Corded Ware) Neolithic phases in the wider region, confirmation for the social patters associated with the Cortaillod culture remains indirect and unproven.

Genomic and isotopic research covering the Middle and Late Neolithic, including the Pfyn culture (c. 3900–3500 BCE), suggest ongoing patterns of patrilocal residence. Studies of ancient genomes from Swiss sites reveal male individuals clustered by kin group over several generations, while females typically came from a broader geographic range. These findings are interpreted as evidence that society at this time was organized such that women wed into an external community while men remained local or within their community. Although less directly addressed than patrilocality, the continuity of kin groups along the male line, as shown by Y-chromosomal data and the presence of father-son-grandson relationships, is strongly indicative of at least partial patrilineal kin organization within these farming groups. [18]

The migratory path between haplogroup G-Z1900 and the haplogroups associated with the beginning of the Muese – Rhine Watershed phylogenetic gap (e.g. G-FGC7516 and G-Z727) followed the general contours of the middle and high portions of the Rhine River as shown in illustration nine.

Illustration Nine: Migratory Path between G-Z1900 and G-Z1817

Effects of the Neolithic Decline

A possible contributing factor to the limited number of subsclades in the Griff(is)(es)(ith) phylogenetic tree is the impact of the Neolithic decline. Between roughly 3450 BCE and 3000 BCE, the population of many Neolithic societies declined throughout western Eurasia due to a combination of factors that are still largely debated. [19] This decline has been referred to as the Neolithic decline.  The Neolithic decline may have occurred during the time when ancestors associated with the G-Z1900 haplogroup lived or impacted the undocumented generations that lived between G-Z1900 and G-Z1817.

Prior to this decline, the Eurasian population increased between 4000 to 3600 BCE, catalyzed by the introduction of agriculture. [20] Other factors associated with the population increase was the spread of technologies such as the production of pottery, the wheel, and animal husbandry practices. [21]

Scientific studies have identified several possible causes for the Neolithic population decline: infectious disease (plague), agricultural crises and environmental degradation, the rise in communicable diseases, nutritional stress and health decline, and social and societal stress. [22]

Recent genomic studies reveal that, unlike surrounding regions that experienced dramatic population replacement and apparent decline, the Meuse-Rhine watershed did not undergo the dramatic Neolithic population collapse seen elsewhere in western Eurasia. Instead, it featured a regionally distinct pattern of demographic persistence and gradual shifts in settlement and subsistence. [23]

While northwest Europe overall saw a marked Neolithic “population crash” associated with agricultural crisis and environmental stress, this did not manifest as a sharp decline in the Meuse-Rhine region. Instead, genetic, archaeological, and ecological evidence suggests a slower, more complex process of change with substantial continuity of indigenous groups until the major influx of Bell Beaker populations.

The Muese-Rhine Watershed in the Neolithic Era

Much of the gentic research associated with the first phase of the ancient DNA European migration between 6500 – 4000 BCE depict Early European Farmers ( EEF), which includes the genetic ancestors of the G haplogroup, migrating from western Anatolia and, in varying degrees, mixing with indigenous Western Hunter-Gatherers (WHG) in Europe. This resulted in a massive genetic turnover of YDNA composition, ranging from 70 to 100 percent, of regional populations. [24] Then a second wave between 3000-2500 BCE of Yamnaya pastoral Western Steppe Herders (WSH), from the Pontic-Caspian region, associated with the Corded Ware cultural complex, spread steppe ancestry into north-central Europe. [25]

The Meuse and Rhine River watersheds played a critical and unique role in shaping Neolithic migration and cultural dynamics in northwestern Europe, particularly through ecological, genetic, and cultural influences. The unique nature of this area may have limited the proliferation of subclades in the Griff(is)(es)(ith) genetic paternal line during this time period.

An intriguing study by Iñigo Olalde and other researchers documented in the wider Rhine-Meuse area an exception to the general pattern European haplogroup migration. The communities in this watershed area existed in the wetlands, riverine areas, and coastal areas of what is currently part of the western and central Netherlands, Belgium and western Germany. [26]

The Rhine-Meuse region showed a unique pattern compared to the rest of continental Europe. Here, a population with approximately fifty percent hunter-gatherer ancestry persisted for up to three millennia longer than elsewhere, lasting from 8500 BCE to 1700 BCE (see illustration ten). This continuity was due to the limited incorporation of Early European Farmer (EEF) ancestry, particularly females, into local communities.

Illustration Ten: Hunter-Gatherer Ancestry Proportions across Europe between 4500-2500 BCE

This prolonged genetic continuity of hunter-gatherer ancestry likely limited diversification within haplogroup G lineages during the ‘first phase or wave’ of migration. Limited Neolithic farmer influx (only about a thirty percent ancestry shift) until the Bell Beaker period reduced external genetic contributions that might have driven subclade formation. This may have contributed to the lack of documented subclades for the Griff(is)(es)(ith) paternal genetic line. [27]

“A striking finding in our study is that the long-term persistence of high hunter-gatherer ancestry was not limited to the core wetlands of the Netherlands, but was also a feature of inland regions of the Rhine (Blatterhöhle cave and Wartberg culture) and Meuse (Belgian Neolithic sites).” [28]

During the ‘second wave’ of migration into the western Netherlands area, the adoption of Corded Ware pottery culture and social practices did not coincide with a significant influx of YDNA R haplogroup steppe ancestry. Individuals in these settlements exhibited Corded Ware Y-chromosome lineages but with minimal steppe genetic input. This contrasts with other parts of Europe, where Corded Ware expansion brought substantial steppe ancestry and changes in haplogroup composition.

Communities in Rhine -Meuse region maintained a semi-agrarian lifestyle combining hunting, fishing, foraging, and limited animal husbandry-crop cultivation. This “extended broad spectrum” subsistence strategy contrasted with the intensive farming of neighboring loess zones [29], reduced the reliance on immigrant agricultural knowledge and allowed the persistence and influence of hunter-gatherer genetic ancestry for 3,000 years longer than in continental Europe. This may have had an tenuating impact on the development of early agrarian communities that were associated with the G haplogroup and the Griff(is)(es)(ith) paternal line. [30]

“(L)arge rivers such as the Scheldt, Meuse and Rhine created a dynamic landscape that included fertile soils favored by farmers, alongside coastlines, beach barriers, river delta wetlands and forested river dunes that continued to support hunting, gathering and fishing practices after 4200 BCE. . . .

“In the Rhine-Meuse area, the wetland communities of the Swifterbant (5th millennium BCE) and Hazendonk cultures (4000-3500 BCE) settled on elevated areas (river and coastal dunes, crevasse splays, and river levees) in a region dominated by water courses and peat bogs. They relied mostly on hunting, gathering, and fishing, but also practiced farming. . . .

“The Swifterbant, Hazendonk and Vlaardingen cultures were all located near water streams, while TRB (Funnelbeaker culture) farmers settled mostly on forested sandy plateaus and their fringes, as did the Michelsberg communities to the south.” [31]

Cultures in the Neolithic Era in the Muese Rhine Watershed

The Rhine-Meuse delta itself was primarily inhabited by the Vlaardingen culture during the Late Neolithic [32] , but the adjacent uplands—especially the Friesian-Drenthian plateau [33] and surrounding sandy regions—were settled by Funnelbeaker communities. These groups established settlements and burial sites distinct from those of the lowland Vlaardingen culture. [34]

Table Six: Cultural Presence in the Meuse-Rhine Watershed in the Neolithic Period

Area	Dominant Culture(s)	Funnelbeaker Evidence
Central/Northern Uplands	Funnelbeaker (TRB)	Settlements, burials, pottery
Rhine-Meuse Delta/Lowlands	Vlaardingen, Hazendonk, Stein Group	Occasional TRB pottery
Southern Uplands	Sparse evidence, Michelsberg	Limited TRB presence

Studies examining the archaeological cultures of the Swifterbant, Hazendonk [35], Vlaardingen, and Funnelbeaker have produced varying degrees of direct evidence that correlate specific haplogroups with these cultures. As of this writing, there are no published, peer-reviewed studies that provide direct, ancient Y-DNA results from individuals ascribed specifically to these Dutch Neolithic and Chalcolithic cultures. Broader Y-DNA studies in the Netherlands (using modern samples) reveal population structure and some continuity, but they do not directly link present-day Y-haplogroups with these archaeological cultures. Some forum and blog discussions mention these cultures in relation to prehistoric Y-DNA, but no direct ancient DNA (aDNA) evidence is available in the scientific literature that conclusively ties a specific haplogroup to Swifterbant, Hazendonk, or Vlaardingen cultures [36]

There are published ancient Y-DNA results that discuss correlations between the Funnelbeaker culture and aDNA haplogroups. Multiple studies, including a summary by Wikipedia and genetic studies referenced there, indicate that most Y-DNA from Funnelbeaker sites belongs to haplogroup I2. A smaller number of samples belong to R1b-V88, Q-FTF30, and G2a. The genetics of this culture show a clear affinity to Neolithic European farmers, with some limited admixture from local hunter-gatherers. [37]

The YDNA genetic ancestors of the Griff(is)(es)(ith) family line may have been part of the Stein Group, Funnelbeaker or Michelsberg cultures. Illustration eleven provides a general depiction of where the Vlaardingen culture and Stein Group culture existed in the delta area and the southern area of the watershed respectively. The Stein group is closely related to the Vlaardingen culture. There is an ongoing debate in Dutch Neolithic archaeology concerning the exact distinction between the Vlaardingen culture and the Stein group, as they are sometimes difficult to differentiate from a typological perspective.  [38]

Illustration Eleven: Distribution of the Vlaardingen Culture and Stein Group

The “Stein culture”, or more specifically, the area and archeological site of Stein, is closely associated with the Linear Pottery culture (LBK), a major archaeological group of the European Neolithic. The LBK flourished from approximately 5500 to 4500 BCE.  The Stein group is mostly located in the Limburg area. Specific excavation reports place the Stein site in Limburg, in the southern Netherlands. Other LBK settlements in the Netherlands have been found elsewhere in South Limburg, including Elsloo. Interestingly, newer discoveries of sites associated with the Vlaardingen Culture, which is culturally linked to the Stein group, are located in the eastern Netherlands, in the area of Nijmegen. [39]

The Griff(is)(es)(ith) ancestors may have lived in areas where the Michelsberg culture existed. The Michelsberg existed in the Neolithic era in parts of Belgium, the Netherlands, and from central Germany to the Paris Basin in northwestern France. The culture developed between 6000 and 4800 years ago, beginning in the Rhineland and Northern France, and is named after the archaeological site of Michelsberg in Baden-Württemberg, Germany. [40]

The Michelsberg people practiced cereal cultivation, but with a reduced crop spectrum compared to earlier Neolithic groups. There was a notable emphasis on stock farming, particularly cattle and pigs, with less reliance on hunting. Organic residue analysis of pottery indicates limited use of ruminant dairy products, suggesting regional variations in diet and food processing. [41]

In the Belgian area, archaeological evidence and scholarly consensus place the existence of the Michelsberg culture approximately between 4400 and 3500 BCE. The Michelsberg culture marks a significant phase in the Neolithic development of Belgium, introducing new technologies and social structures that influenced subsequent cultures in the region. [42]

Michelsberg communities constructed ditched enclosures, sometimes on hilltops, which may have served as communal, defensive, or ceremonial spaces. Archaeological finds include well-preserved domed ovens which may indicate communal food processing or adaptation to local climate conditions. Evidence suggests both enclosed and open settlements, with some sites showing rows of small houses or sunken-floor buildings. [43]

The widespread distribution of flint tools and axes among the Michelsberg sites, as well as the presence of exotic materials like jade and variscite, points to extensive trade networks connecting areas presently known as Belgium with regions as far as modern day Italy and Spain. [44]

There is currently no direct archaeological or anthropological evidence specifically indicating patrilocality (post-marital residence with the husband’s group) or patrilineal (descent and inheritance through the male line) practices in the Vlaardingen, Hazendonk, or Stein Group cultures in the Rhine–Meuse–Scheldt delta region during the Neolithic period. Without DNA kinship analyses, specific burial clustering, or other direct archaeological markers, any statements about their kinship or residence patterns remain speculative and unsupported by present data. [45]

Another cultural group that may have been associated with the haplogroups is the Funnelbeaker culture. Archaeological evidence confirms that the Funnelbeaker culture was present in the uplands of the central, northern, and eastern Netherlands, which are part of the Rhine and Meuse watershed. These areas were characterized by sandy plateaus and higher grounds suitable for early farming and settlement.

Illustration twelve depicts some level of contact or exchange between the Vlaadingen, Funnelbeaker and Stein cultures in the watershed area based on archeaological sites. The Funnelbeaker culture is associated with megalithic tombs and collective burials, which are found in the upland regions north of the Rhine and Meuse rivers, further supporting their presence in the broader watershed. [46]

Illustration Twelve: The Cultural Landscape of the Netherlands 3400 – 2850 BCE

A study conducted by Julia Menne and Mirco Brunner investigated the transition from the late Swifterbant culture to the Funnelbeaker Westgroup (TRB) culture, an early cultural manifestation of the Early European Farmers (EEF), in northwestern Europe. They focused on the shift from Mesolithic hunter-gatherer societies to fully Neolithic communities. (See illustration thirteen)

Illustration Thirteen: Transition from and Relationship Between Hunter Gatherer (Swifterbant) and Early Farmer (Funnelbeaker) Cultural Sites

The study area covers northwestern Germany and the north/northeast Netherlands, spanning from 6000 to 2600 (radio-carbon calibrated) BCE. Their goal was to clarify whether there were cultural discontinuities (hiatuses) or gradual transitions between these archaeological phases. Their study points to evidence of overlap and continuity between the transition from the Late Swifterbant sites to Funnelbeaker culture. The transition to the Neolithic era in this region was a ‘mosaic process’, involving both coexistence and transition between the two groups. [47]

The Funnelbeaker culture along with hunter gatherer cultures were precursor cultures to the Corded Ware culture in the Rhine-Meuse watershed area, both chronologically and in terms of population continuity. However, rather than a simple linear succession, the transition involved a mix of cultural replacement, intermarriage, and local adaptation, with the Rhine-Meuse region showing particularly strong evidence for local cultural persistence and a gradual incorporation of Corded Ware cultural elements. [48]

There is strong evidence that the Funnelbeaker culture (also known as TRB) practiced both patrilocality and patrilineal kinship practices. Genetic analysis of individuals buried in Funnelbeaker megalithic tombs, such as the Frälsegården passage grave in Sweden, demonstrates that the vast majority of males in these graves were descended from a single male ancestor. In contrast, the women buried alongside them were predominantly unrelated, suggesting they had married into the group. This pattern is a hallmark of patrilineal descent, where inheritance and group membership are traced through the male line. [49]

Studies of mitochondrial DNA and Y-chromosome DNA from Funnelbeaker graves show higher mitochondrial diversity compared to Y-chromosome diversity. This indicates that women frequently came from outside the local group—supporting the idea that, after marriage, women moved to live with their husband’s group (patrilocality). [50]

The presence of collective burials containing related males, as well as the structure of passage graves and dolmens [51] , further supports a patrilineal and patrilocal framework in the Funnelbeaker culture. This social organization is consistent with other Middle Neolithic European farming populations, where collective burials often comprised males related by descent and unrelated incoming females.

The Funnelbeaker culture dominated much of north-central Europe, including the Rhine-Meuse region, from roughly 4300–2800 BCE. The Corded Ware culture appeared in the early third millennium BCE (around 2900–2650 BCE), eventually replacing the Funnelbeaker culture in this area. [52] (See table seven)

Table Seven: Funnelbeaker vs. Corded Ware in the Rhine-Meuse Watershed

Aspect	Funnelbeaker Culture (TRB)	Corded Ware Culture (CWC)
Timeframe	~4300–2800 BCE	~2900–2300 BCE
Population ancestry	Early European Farmer + hunter-gatherer	Steppe ancestry (male), EEF (female)
Cultural transition	Gradually replaced by CWC	Adopted some TRB practices, reused sites
Genetic interaction	Maternal lines persist in CWC	Intermarriage with local TRB women
Regional dynamics	Strong local continuity in Rhine-Meuse	Initial low steppe ancestry in region

In some regions, the transition from Funnelbeaker to Corded Ware showed signs of continuity, with Corded Ware adopting and adapting local practices and even reusing Funnelbeaker megalithic tombs. [53] Genetic studies indicate that women from the Funnelbeaker culture were incorporated into the Corded Ware population, suggesting intermarriage and some degree of population continuity, especially on the maternal side. [54]

The two cultures coexisted for several centuries in the Rhine-Meuse area, with indigenous Funnelbeaker communities maintaining their practices even after the arrival of Corded Ware groups. Rather than a rapid or violent replacement, the transition involved ongoing interaction and gradual blending of cultural elements. [55]

Source:

Feature Banner: The banner at the top of the story features a map of the phylogenetic gaps discussed in the story. The maps was generated by taking a snapshop from the FamilyTreeDNA Globetrekker^TM video of the migratory path of my YDNA descendants over time. The map shows the migratory path of selected most common recent ancestors and their respective estimated dates of birth. Another map in the banner depicts the Muesse and Rhine River watershed that is associated with this phylogenetic gap. . In addition, various cultures and features associated with time periods within this period of time are depicted.

[1] Ingram, Owen, What is Ecological Fallacy – Causes & Examples, Rev 5 Jan 2025, ResearchProspect, https://www.researchprospect.com/what-is-ecological-fallacy/

Nikolopoulou, Kassiani, What Is Ecological Fallacy? | Definition & Example, 7 Jan 2023, Scribbr, https://www.scribbr.com/fallacies/ecological-fallacy/

For a practical examples, see:

K.N. Lala, & M.W. Feldman, Genes, culture, and scientific racism, Proc. Natl. Acad. Sci. U.S.A. 121 (48) e2322874121,https://doi.org/10.1073/pnas.2322874121 (2024).

Foley RA, Lahr MM. The evolution of the diversity of cultures. Philos Trans R Soc Lond B Biol Sci. 2011 Apr 12;366(1567):1080-9. doi: 10.1098/rstb.2010.0370. PMID: 21357230; PMCID: PMC3049104 PubMed https://pmc.ncbi.nlm.nih.gov/articles/PMC3049104/

Aaron P Ragsdale, Kevin R Thornton, Multiple Sources of Uncertainty Confound Inference of Historical Human Generation Times, Molecular Biology and Evolution, Volume 40, Issue 8, August 2023, msad160, https://doi.org/10.1093/molbev/msad160

[2] Linear Pottery Culture (jan 1, 5500 BC – jan 1, 4500 BC), Public TimeLines, Time Graphics, https://time.graphics/period/3613007

Linear Pottery culture, Wikipedia, This page was last edited on 3 April 2025, https://en.wikipedia.org/wiki/Linear_Pottery_culture

Danubian Culture, Wikipedia, This page was last edited on 15 June 2024, https://en.wikipedia.org/wiki/Danubian_culture

[3] Linear Pottery culture, Wikipedia, This page was last edited on 3 April 2025,, https://en.wikipedia.org/wiki/Linear_Pottery_culture

Danubian Culture, Wikipedia, This page was last edited on 15 June 2024, https://en.wikipedia.org/wiki/Danubian_culture

[4] Linear Pottery Culture, Wikipedia, This page was last edited on 18 July 2025, https://en.wikipedia.org/wiki/Linear_Pottery_culture

Dolukhanov, Pavel; Shukurov, A; Gronenborn, D; Sokoloff, D; Timofeev, V; Zaitseva, G; et al. (2005). “The Chronology of Neolithic Dispersal in Central and Eastern Europe” (PDF). Journal of Archaeological Science. 32 (10): 1441–1458. Bibcode:2005JArSc..32.1441D. doi:10.1016/j.jas.2005.03.02

Dolukhanov, Pavel; Shukurov, Anvar (2003). “Modelling the Neolithic Dispersal in Northern Eurasia” , Documenta Praehistorica XXXI. Department of Archaeology, Faculty of Arts, University of Ljubljana. Archived from the original (PDF) on 27 September 2007, https://web.archive.org/web/20070927045407/http://arheologija.ff.uni-lj.si/documenta/pdf31/31dolukhanov.pdf

[5] Linear Pottery Culture (Jan 1, 5500 BC – Jan 1, 4500 BC), Public TimeLines, Time Graphics, https://time.graphics/period/3613007

Linear Pottery culture, Wikipedia, This page was last edited on 3 April 2025,, https://en.wikipedia.org/wiki/Linear_Pottery_culture

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David M, An Introduction to the Neolithic Linearbandkeramik Culture, 6 Dec 2013,These Bones of Mine, https://thesebonesofmine.wordpress.com/2013/12/06/an-introduction-to-the-neolithic-linearbandkeramik-culture/

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[6] Linear Pottery culture, Wikipedia, This page was last edited on 3 April 2025,, https://en.wikipedia.org/wiki/Linear_Pottery_culture

[7] The Tyrolean Alps are a mountain range located primarily in the Tyrol region of Austria and the South Tyrol region of Italy. They form part of the larger Eastern Alps.

Griff(is)(es)(ith) Haplogroups in Context of the Tyrolean Alps

Tyrol, Wikipedia, This page was last edited on 3 July 2025, https://en.wikipedia.org/wiki/Tyrol

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[8] The spread of G-L497 can be traced through several major population movements:

Regional Expansion: G-L497 lineages are also tied to the Etruscans and Germanic groups such as the Goths and Franks. Notable hotspots today include the Tyrolean Alps, Switzerland, Northern Italy, southwestern Germany, and eastern France

Neolithic Migration: Early agriculturalists carrying G-L497 expanded from Anatolia into Europe, mainly via the Balkans and up the Danube River into Central Europe. This established the haplogroup in areas like Germany, Austria, Switzerland, and Northern Italy.

Bronze Age Dispersal: During the Bronze Age (c. 1800 BCE), G-L497 moved from Italy northwards into Central and Northern Europe.

Celtic Expansion: The haplogroup is associated with the Hallstatt and La Tène Celtic cultures, which further spread G-L497 throughout the Alpine regions, France, and into the British Isles. Elite burials from the Hallstatt period have yielded G-L497, indicating its prominence among early Celts.

G-L497 is closely linked to the following historical cultures::

• Neolithic and Chalcolithic agricultural societies;
• Hallstatt and La Tène Celts;
• Italic tribes and the Etruscans in Italy; and
• Early Germanic tribes, such as the Suebi and Franks, especially in southern Germany and the Rhine region.

Hay, Maciamo, New map of Y-gaplogroup G2a-L497, 22 May 2017, Eupedia, https://www.eupedia.com/forum/threads/new-map-of-y-gaplogroup-g2a-l497.34104/

G-L497 Y-DNA Project, FamilyTreeDNA, https://www.familytreedna.com/groups/g-ydna/about

Rootsi S, Myres NM, Lin AA, Järve M, King RJ, Kutuev I, Cabrera VM, Khusnutdinova EK, Varendi K, Sahakyan H, Behar DM, Khusainova R, Balanovsky O, Balanovska E, Rudan P, Yepiskoposyan L, Bahmanimehr A, Farjadian S, Kushniarevich A, Herrera RJ, Grugni V, Battaglia V, Nici C, Crobu F, Karachanak S, Hooshiar Kashani B, Houshmand M, Sanati MH, Toncheva D, Lisa A, Semino O, Chiaroni J, Di Cristofaro J, Villems R, Kivisild T, Underhill PA. Distinguishing the co-ancestries of haplogroup G Y-chromosomes in the populations of Europe and the Caucasus. Eur J Hum Genet. 2012 Dec;20(12):1275-82. doi: 10.1038/ejhg.2012.86. Epub 2012 May 16. PMID: 22588667; PMCID: PMC3499744. (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC3499744/

Berger B, Niederstätter H, Erhart D, Gassner C, Schennach H, Parson W. Reprint of: high resolution mapping of Y haplogroup G in Tyrol (Austria). Forensic Sci Int Genet. 2013 Dec;7(6):624-631. doi: 10.1016/j.fsigen.2013.09.006. Epub 2013 Sep 27. Erratum in: Forensic Sci Int Genet. 2014 Mar;9:191. PMID: 24099688. (PubMed) https://pubmed.ncbi.nlm.nih.gov/24099688/ see also https://www.researchgate.net/publication/291505390_High_resolution_mapping_of_Y_haplogroup_G_in_Tyrol_Austria_Reprinted_from_Forensic_Science_International_Genetics_vol_7_pg_529-536_2013

[9] G-L497 Y-DNA Project, FamilyTreeDNA, https://www.familytreedna.com/groups/g-ydna/about

Rootsi S, et al., Distinguishing the co-ancestries of haplogroup G Y-chromosomes in the populations of Europe and the Caucasus. Eur J Hum Genet. 2012 Dec;20(12):1275-82. doi: 10.1038/ejhg.2012.86. Epub 2012 May 16. PMID: 22588667; PMCID: PMC3499744. (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC3499744/

Hay, Maciamo, New map of Y-gaplogroup G2a-L497, 22 May 2017, Eupedia, https://www.eupedia.com/forum/threads/new-map-of-y-gaplogroup-g2a-l497.34104/

[10] Berger B, et al., Reprint of: high resolution mapping of Y haplogroup G in Tyrol (Austria). Forensic Sci Int Genet. 2013 Dec;7(6):624-631. doi: 10.1016/j.fsigen.2013.09.006. Epub 2013 Sep 27. Erratum in: Forensic Sci Int Genet. 2014 Mar;9:191. PMID: 24099688. (PubMed) https://pubmed.ncbi.nlm.nih.gov/24099688/

Hay, Maciamo, Haplogroup G2a (Y-DNA), Jul 2023, Eupedia, https://www.eupedia.com/europe/Haplogroup_G2a_Y-DNA.shtml

[11] Andreas Putzer, Daniela Festi, Sophie Edlmair , Klaus Oeggl, The development of human activity in the high altitudes of the Schnals Valley (South Tyrol/Italy) from the Mesolithic to modern periods, Journal of Archeological Science: Reports, 21 Jan 2016, 6, 136-147, https://www.academia.edu/30556244/The_development_of_human_activity_in_the_high_altitudes_of_the_Schnals_Valley_South_Tyrol_Italy_from_the_Mesolithic_to_modern_periods

History of Tyrol, Wikipedia, This page was last edited on 16 February 2025, https://en.wikipedia.org/wiki/History_of_Tyrol

Neolithic Italy, Wikipedia, This page was last edited on 23 March 2025, https://en.wikipedia.org/wiki/Neolithic_Italy

Stevenson, Joan C., Neolithic Age Europe, 2022, EBSCO, https://www.ebsco.com/research-starters/anthropology/neolithic-age-europe

Mathieson I, Alpaslan-Roodenberg S, Posth C, Szécsényi-Nagy A, Rohland N, Mallick S, Olalde I, Broomandkhoshbacht N, Candilio F, Cheronet O, Fernandes D, Ferry M, Gamarra B, Fortes GG, Haak W, Harney E, Jones E, Keating D, Krause-Kyora B, Kucukkalipci I, Michel M, Mittnik A, Nägele K, Novak M, Oppenheimer J, Patterson N, Pfrengle S, Sirak K, Stewardson K, Vai S, Alexandrov S, Alt KW, Andreescu R, Antonović D, Ash A, Atanassova N, Bacvarov K, Gusztáv MB, Bocherens H, Bolus M, Boroneanţ A, Boyadzhiev Y, Budnik A, Burmaz J, Chohadzhiev S, Conard NJ, Cottiaux R, Čuka M, Cupillard C, Drucker DG, Elenski N, Francken M, Galabova B, Ganetsovski G, Gély B, Hajdu T, Handzhyiska V, Harvati K, Higham T, Iliev S, Janković I, Karavanić I, Kennett DJ, Komšo D, Kozak A, Labuda D, Lari M, Lazar C, Leppek M, Leshtakov K, Vetro DL, Los D, Lozanov I, Malina M, Martini F, McSweeney K, Meller H, Menđušić M, Mirea P, Moiseyev V, Petrova V, Price TD, Simalcsik A, Sineo L, Šlaus M, Slavchev V, Stanev P, Starović A, Szeniczey T, Talamo S, Teschler-Nicola M, Thevenet C, Valchev I, Valentin F, Vasilyev S, Veljanovska F, Venelinova S, Veselovskaya E, Viola B, Virag C, Zaninović J, Zäuner S, Stockhammer PW, Catalano G, Krauß R, Caramelli D, Zariņa G, Gaydarska B, Lillie M, Nikitin AG, Potekhina I, Papathanasiou A, Borić D, Bonsall C, Krause J, Pinhasi R, Reich D. The genomic history of southeastern Europe. Nature. 2018 Mar 8;555(7695):197-203. doi: 10.1038/nature25778. Epub 2018 Feb 21. PMID: 29466330; PMCID: PMC6091220. (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC6091220/

Neolithic, Wikipedia, This page was last edited on 22 July 2025, https://en.wikipedia.org/wiki/Neolithic

6th millennium BC, Wikipedia, This page was last edited on 22 July 2025, https://en.wikipedia.org/wiki/6th_millennium_BC

Detlef Gronenborn, Migration, acculturation and culture change in western temperate Eurasia, 6500–5000 cal BC, Documenta Praehistorica XXX, 79-91, https://pdfs.semanticscholar.org/4c78/ed1587f5f471fd5209b8fedfd7ec42ec6e56.pdf

[12] Neolithic Europe, Wikipedia, This page was last edited on 19 July 2025, https://en.wikipedia.org/wiki/Neolithic_Europe

See also:

Swiss Pile Dwellings, Milwaulkee Public Museum, https://www.mpm.edu/research-collections/anthropology/online-collections-research/robenhausen-site/culture

Cortaillod culture, Wikipeda, This page was last edited on 27 May 2025, https://en.wikipedia.org/wiki/Cortaillod_culture

Pfyn – Pre-Roman Era in German (http://www.hls-dhs-dss.ch/textes/d/D1982.php), French (http://www.hls-dhs-dss.ch/textes/f/F1982.php) and Italian (http://www.hls-dhs-dss.ch/textes/i/I1982.php) in the online Historical Dictionary of Switzerland

Francesco Menotti, Living on the lake in prehistoric Europe: 150 years of lake-dwelling research (https://books.google.com/books?id=OEK5PyQUtegC&pg=PA154). Routledge, 2004, pg. 57.

Culture | Milwaukee Public Museum. (n.d.). https://www.mpm.edu/research-collections/anthropology/online-collections-research/robenhausen-site/culture

Rainer Berger; Hans Eduard Suess (1979). Radiocarbon dating: proceedings of the ninth international conference, Los Angeles and La Jolla, 1976 (https://books.google.com/books?id=YupA6DrPQJIC&pg=PA104 ). University of California Press. pg. 104–107.

Schibler, J. 2006. “The economy and environment of the 4th and 3rd millennia BC in the northern Alpine foreland based on studies of animal bones”. Environmental Archaeology 11(1):
49-65

[13] Swiss Pile Dwellings, Milwaulkee Public Museum, https://www.mpm.edu/research-collections/anthropology/online-collections-research/robenhausen-site/culture

[14] Neolithic Europe, Wikipedia, This page was last edited on 19 July 2025, https://en.wikipedia.org/wiki/Neolithic_Europe

Swiss Pile Dwellings, Milwaulkee Public Museum, https://www.mpm.edu/research-collections/anthropology/online-collections-research/robenhausen-site/culture

[15] Prehistoric pile dwellings around Lake Zurich, Wikipedia, This page was last edited on 27 April 2025, https://en.wikipedia.org/wiki/Prehistoric_pile_dwellings_around_Lake_Zurich

Prehistoric Pile Dwellings around the Alps, Experience Hauterive, https://www.myswitzerland.com/en-ch/experiences/pile-dwellings-around-the-alps/

[16] David M. , An Introduction to the Neolithic Linearbandkeramik Culture, 6 Dec, 2013, These Bones of Mine, https://thesebonesofmine.wordpress.com/2013/12/06/an-introduction-to-the-neolithic-linearbandkeramik-culture/

[17] Ibid

[18] Furtwängler, A., Rohrlach, A.B., Lamnidis, T.C. et al. Ancient genomes reveal social and genetic structure of Late Neolithic Switzerland. Nat Commun 11, 1915 (2020). https://doi.org/10.1038/s41467-020-15560-x

Furtwängler, A., et al , Neolithic Genomes From Modern-Day Switzerland Indicate Parallel Ancient Societies, 20 Apr 2020, Max Planck Institute of Geoanthropology, https://www.shh.mpg.de/1671901/neolithic-genomes-from-modern-day-switzerland

[19] Rascovan, Nicolas; Sjögren, Karl-Göran; Kristiansen, Kristian; Nielsen, Rasmus; Willerslev, Eske; Desnues, Cristelle; Rasmussen, Simon (10 January 2019), “Emergence and Spread of Basal Lineages of Yersinia pestis during the Neolithic Decline”. Cell. 176(2): 295–305. doi:10.1016/j.cell.2018.11.005. PMID 30528431. S2CID 54447284 https://www.cell.com/cell/fulltext/S0092-8674(18)31464-8

Zhang, Sarah (6 December 2018). “An Ancient Case of the Plague Could Rewrite History”. The Atlantic. Retrieved 13 November 2019. https://www.theatlantic.com/science/archive/2018/12/4900-year-old-case-plague-sweden/577315/

[20] Zhang, Sarah (6 December 2018). “An Ancient Case of the Plague Could Rewrite History”. The Atlantic. Retrieved 13 November 2019. https://www.theatlantic.com/science/archive/2018/12/4900-year-old-case-plague-sweden/577315/

Colledge, Sue; Conolly, James; Crema, Enrico; Shennan, Stephen (2019-01-10). “Neolithic population crash in northwest Europe associated with agricultural crisis”. Quaternary Research. 92 (3): 686–707. Bibcode:2019QuRes..92..686C. doi:10.1017/qua.2019.42. ISSN 0033-5894. S2CID 202186375, https://www.cambridge.org/core/journals/quaternary-research/article/abs/neolithic-population-crash-in-northwest-europe-associated-with-agricultural-crisis/11F405EB351C8A86020C17A647C1C001

[21] Rascovan, Nicolas; et al., (10 January 2019), “Emergence and Spread of Basal Lineages of Yersinia pestis during the Neolithic Decline”. Cell. 176(2): 295–305. doi:10.1016/j.cell.2018.11.005. PMID 30528431. S2CID 54447284 https://www.cell.com/cell/fulltext/S0092-8674(18)31464-8

[22] The following are the primary causes proposed and supported by recent archaeological and genetic research:

Infectious Disease (Plague): Recent ancient DNA analyses reveal that ancestral forms of the plague bacterium (Yersinia pestis) were present among Neolithic populations in Europe, especially in Scandinavia and Northwestern Europe. These studies suggest that repeated plague epidemics could have caused drastic reductions in population size, in a manner similar to the later Black Death epidemics. This finding has shifted considerable support toward disease, particularly plague,as a major factor in the Neolithic decline, although it does not exclude other contributory factors.

Seersholm, F.V., Sjögren, KG., Koelman, J. et al. Repeated plague infections across six generations of Neolithic Farmers. Nature 632, 114–121 (2024). https://doi.org/10.1038/s41586-024-07651-2

Univeristy of Copenhagen, DNA analyses show the plague may have caused the downfall of Stone Age farmers, 10 Jul 2024, , Phys.Org, https://phys.org/news/2024-07-dna-analyses-plague-downfall-stone.html

Molinek, Rudy, Did Plague Cause the Mysterious Collapse of Europe’s Early Farmers 5,000 Years Ago?, Smithsonian Magazine, https://www.smithsonianmag.com/smart-news/did-plague-cause-the-mysterious-collapse-of-europes-early-farmers-5000-years-ago-180984688/

Agricultural Crisis and Environmental Degradation: The Neolithic decline is also associated with evidence of worsening agricultural conditions. Environmental factors such as declining cereal yields, soil exhaustion from intensive agriculture, and possibly adverse climate changes are linked to the abandonment of settlements and regrowth of forests in formerly cultivated areas. These factors could have resulted in food shortages, malnutrition, and increased vulnerability to disease.

Neolithic Decline, Wikipedia, This page was last edited on 24 June 2025, https://en.wikipedia.org/wiki/Neolithic_decline#cite_note-atlantic-3

Sean S. Downey, W. Randall Haas Jr., and Stephen J. Shennan, European Neolithic societies showed early warning signals of population collapse, August 29, 2016, PNAS, 113 (35) 9751-9756, https://www.pnas.org/doi/10.1073/pnas.1602504113

Colledge S, Conolly J, Crema E, Shennan S. Neolithic population crash in northwest Europe associated with agricultural crisis. Quaternary Research. 2019;92(3):686-707. doi:10.1017/qua.2019.42 https://www.cambridge.org/core/journals/quaternary-research/article/abs/neolithic-population-crash-in-northwest-europe-associated-with-agricultural-crisis/11F405EB351C8A86020C17A647C1C001

Rise in Communicable Diseases: The transition to settled, farming-based societies led to higher population densities and close cohabitation with domesticated animals. This new lifestyle increased the risk of zoonotic (animal-derived) and waterborne diseases, elevating general mortality rates alongside disease outbreaks.

Neolithic demographic transition, Wikipedia, This page was last edited on 28 May 2025, https://en.wikipedia.org/wiki/Neolithic_demographic_transition

Marko Porčić, Tamara Blagojević, Jugoslav Pendić and Sofija Stefanović, The Neolithic Demographic Transition in the Central Balkans: population dynamics reconstruction based on new radiocarbon evidence, 30 November 2020, Philosophical Transations of the Royal Society B Biological Sciences, https://royalsocietypublishing.org/doi/10.1098/rstb.2019.0712

Bocquet-Appel JP. Explaining the Neolithic Demographic Transition. The Neolithic Demographic Transition and its Consequences. 2008:35–55. doi: 10.1007/978-1-4020-8539-0_3. PMCID: PMC7120555. (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC7120555/

Nutritional Stress and Health Decline: The shift from diverse wild food sources to a carbohydrate-heavy agricultural diet, often lacking in essential nutrients, may have led to poorer health, contributing to increased mortality from both malnutrition and infectious disease.

Bocquet-Appel JP. Explaining the Neolithic Demographic Transition. The Neolithic Demographic Transition and its Consequences. 2008:35–55. doi: 10.1007/978-1-4020-8539-0_3. PMCID: PMC7120555. (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC7120555/

Social and Societal Stress: Archaeological evidence hints at possible societal destabilization, including population pressures, conflict, and the collapse of traditional cultural practices (such as the cessation of megalith building), potentially exacerbating demographic decline.

Neolithic Decline, Wikipedia, This page was last edited on 24 June 2025, https://en.wikipedia.org/wiki/Neolithic_decline#cite_note-atlantic-3

Sean S. Downey, W. Randall Haas Jr., and Stephen J. Shennan, European Neolithic societies showed early warning signals of population collapse, August 29, 2016, PNAS, 113 (35) 9751-9756, https://www.pnas.org/doi/10.1073/pnas.1602504113

Adrian Timpson, Sue Colledge, Enrico Crema, Kevan Edinborough, Tim Kerig, Katie Manning, Mark G. Thomas, Stephen Shennan, Reconstructing regional population fluctuations in the European Neolithic using radiocarbon dates: a new case-study using an improved method, Journal of Archaeological Science, Volume 52, 2014, Pages 549-557, ISSN 0305-4403, https://doi.org/10.1016/j.jas.2014.08.011.
(https://www.sciencedirect.com/science/article/pii/S0305440314002982 )

[23] Iñigo Olalde, et al, Long-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups, bioRxiv, 25 Mar 2025, page 19, doi: https://doi.org/10.1101/2025.03.24.644985 , https://www.biorxiv.org/content/10.1101/2025.03.24.644985v1.full.pdf

[24] See for example:

Curry, Andrew, The First Europeans Weren’t Who Your Might Think, National Geographic Magazine, August 2019, online: https://www.nationalgeographic.com/culture/article/first-europeans-immigrants-genetic-testing-feature

Reich, David (2018). Who We are and how We Got Here: Ancient DNA and the New Science of the Human Past. Oxford University Press.

Haak, W. et al. Massive migration from the steppe was a source for Indo-European languages in Europe. Nature 522, 207–211 (2015).

Allentoft, M. E. et al. Population genomics of Bronze Age Eurasia. Nature 522, 167–172 (2015). 

Olalde, I. et al. The Beaker phenomenon and the genomic transformation  of northwest Europe. Nature 555, 190–196 (2018).

Papac, L. et al. Dynamic changes in genomic and social structures in third  millennium BCE central Europe. Sci. Adv. 7, (2021).

Brandt, G. et al. Ancient DNA reveals key stages in the formation of Central European mitochondrial genetic diversity. Science 342, 257–261 (2013).

[25] Iñigo Olalde, et al, Long-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups, bioRxiv, 25 Mar 2025, doi: https://doi.org/10.1101/2025.03.24.644985 , https://www.biorxiv.org/content/10.1101/2025.03.24.644985v1.full.pdf

Olalde I, et al , Long-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups. bioRxiv [Preprint]. 2025 Mar 25:2025.03.24.644985. doi: 10.1101/2025.03.24.644985. PMID: 40196638; PMCID: PMC11974744 (PubMed) https://pubmed.ncbi.nlm.nih.gov/40196638/

Fernández-Götz, Manuel, and others (eds), Rethinking Migrations in Late Prehistoric Eurasia (London, 2022; online edn, British Academy Scholarship Online, 18 May 2023), https://doi.org/10.5871/bacad/9780197267356.001.0001

Khan, Razib, Facing Facts, even fraught ones: the quest for proto-Indo-Europeans in 2023: How genetics illuminates the rise of Eurasian steppe pastoralism, 23 Sep 2023, Unsupervised Learning, https://www.razibkhan.com/p/facing-facts-even-fraught-ones-the

[26] Iñigo Olalde, et al, Long-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups, bioRxiv, 25 Mar 2025, doi: https://doi.org/10.1101/2025.03.24.644985 , https://www.biorxiv.org/content/10.1101/2025.03.24.644985v1.full.pdf

Allentoft ME et al., Population genomics of Bronze Age Eurasia. Nature 2015; 522: 167–172. https://doi.org/10.1038/nature14507 PMID: 26062507

HaakWet al., Massive migration from the steppe was a source for Indo-European languages in Europe. Nature 2015; 522: 207–211. https://doi.org/10.1038/nature14317 PMID: 25731166

Olalde I et al., The Beaker Phenomenon and the Genomic Transformation of Northwest Europe. Nature 2018; 555: 190–196. https://doi.org/10.1038/nature25738 PMID: 29466337

[27] Iñigo Olalde, et al, Long-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups, bioRxiv, 25 Mar 2025, doi: https://doi.org/10.1101/2025.03.24.644985 , https://www.biorxiv.org/content/10.1101/2025.03.24.644985v1.full.pdf

Olalde I, et al , Long-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups. bioRxiv [Preprint]. 2025 Mar 25:2025.03.24.644985. doi: 10.1101/2025.03.24.644985. PMID: 40196638; PMCID: PMC11974744 (PubMed) https://pubmed.ncbi.nlm.nih.gov/40196638/

[28] Iñigo Olalde, et al, Long-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups, bioRxiv, 25 Mar 2025, Page 2-3, doi: https://doi.org/10.1101/2025.03.24.644985 , https://www.biorxiv.org/content/10.1101/2025.03.24.644985v1.full.pdf

[29] Loess tends to develop into very rich soils. Under appropriate climatic conditions, it is some of the most agriculturally productive terrain in the world.

Loess zones, also known as loess plains or loess hills, are areas characterized by extensive deposits of wind-blown silt called loess. These deposits are typically yellowish-gray in color and can be quite thick, forming relatively flat or gently rolling landscapes

Getis, Arthur; Judith Getis and Jerome D. Fellmann (2000). Introduction to Geography, Seventh Edition. Boston: McGraw Hill. p. 99 https://archive.org/details/introductiontoge00geti

[30] Iñigo Olalde et al, Long-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups, bioRxiv, 25 Mar 2025, doi: https://doi.org/10.1101/2025.03.24.644985 , https://www.biorxiv.org/content/10.1101/2025.03.24.644985v1.full.pdf

Olalde I, et al, Long-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups. bioRxiv [Preprint]. 2025 Mar 25:2025.03.24.644985. doi: 10.1101/2025.03.24.644985. PMID: 40196638; PMCID: PMC11974744 (PubMed) https://pubmed.ncbi.nlm.nih.gov/40196638/

[31] Quote:

Iñigo Olalde et al, Long-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups, bioRxiv, 25 Mar 2025, doi: https://doi.org/10.1101/2025.03.24.644985 , https://www.biorxiv.org/content/10.1101/2025.03.24.644985v1.full.pdf, Page 2-3

See also:

Cohen, Kim, Sediment History, Utrecht University, https://www.uu.nl/en/research/water-climate-and-future-deltas/storylines/sediments-matter/follow-the-sediment/sediment-history

[32] “This hunting and fishing culture was sedentary and semi-nomadic. In the old dunes at the then much further east lying coastline the remains of some peasant settlements were found. From these findings it can be deduced that animal husbandry (sheep and goats) and agriculture (wheat and barley) had been practised on a small scale. Vlaardingen culture also had traits of a Mesolithichunter-gatherer society because agriculture in much of the Wadden Sea area and the Rhine–Meuse–Scheldt delta was difficult. Hunting and fishing were important means of livelihood.“

Vlaardingen culture, Wikipedia, This page was last edited on 21 June 2025, https://en.wikipedia.org/wiki/Vlaardingen_culture

See also:

Brinkkemper, Otto, Erik Drenth, Jørn Zeiler, An outline of the subsistence of the Vlaardingen culture from the Netherlands Révue Archéologique de Picardi, 1 Jan 2011, Reviews in Aquaculture Spécial 28:207-220, 10.3406/pica.2011.3331, https://www.researchgate.net/publication/248391480_An_outline_of_the_subsistence_of_the_Vlaardingen_culture_from_the_Netherlands_Revue_Archeologique_de_Picardi or https://www.persee.fr/doc/pica_1272-6117_2011_hos_28_1_3331

Welmoed, A. Out, Walter Dörfler, The best of both worlds: Human impact and plant subsistence at the Middle and Late Neolithic semi-agricultural site of Hekelingen III (2900–2500 BC), Quaternary International, Vol 426, Part B, May 2017, 41-63, https://www.sciencedirect.com/science/article/abs/pii/S1040618215012070

Prehistory of the Netherlands, Wikipedia, This page was last edited on 9 July 2025, https://en.wikipedia.org/wiki/Prehistory_of_the_Netherlands

[33] The Friesian-Drenthe Plateau is a landscape feature in the northern Netherlands, encompassing large parts of the provinces of Drenthe and Friesland, the northwestern part of the province of Overijssel, and a small part of Groningen. This plateau has existed since the Pleistocene era. 

Fokkens, Harry, Drowned Landscape, The Occupation of the Western Part of the Frisian-Drentian Plateau, 4400BC – AD 500, Netherlands: Assen: Van Gorcum & Comp, 1998 https://scholarlypublications.universiteitleiden.nl/handle/1887/11705

M.J.L.Th. Niekus, R.T. van Balen, J.M.G. Bongers, J.H.A. Bosch, D.C. Brinkhuizen, H. Huisman, J. Jelsma, L. Johansen, C. Kasse, S.J. Kluiving, H. Paas, E. Schrijer, D. Stapert, News from the north: A late Middle Palaeolithic site rich in handaxes on the Drenthe–Frisian till plateau near Assen, the Netherlands; first results of a trial excavation, Quaternary International, Volume 411, Part A, 2016, Pages 284-304, ISSN 1040-6182,
https://doi.org/10.1016/j.quaint.2015.06.034
(https://www.sciencedirect.com/science/article/pii/S1040618215006345 )

[34] Folkens, Harry, Background to Dutch Beakers. A Critical Review of the Dutch Model, Pages 9 – 36 in Harry Fokkens and Franco Nicolis, eds, Background to Beakers, van Woerdekom: Sidestone Press, 2012, https://www.sidestone.com/openaccess/9789088900846.pdf

Fokkens, Harry, Drowned Landscape, The Occupation of the Western Part of the Frisian-Drentian Plateau, 4400BC – AD 500, Netherlands: Assen: Van Gorcum & Comp, 1998 https://scholarlypublications.universiteitleiden.nl/handle/1887/11705

Funnelbeaker culture, Wikipedia, This page was last edited on 2 July 2025, https://en.wikipedia.org/wiki/Funnelbeaker_culture

Corded Ware culture, Wikipedia, This page was last edited on 23 June 2025, https://en.wikipedia.org/wiki/Corded_Ware_culture

Olalde I, et al , Long-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups. bioRxiv [Preprint]. 2025 Mar 25:2025.03.24.644985. doi: 10.1101/2025.03.24.644985. PMID: 40196638; PMCID: PMC11974744 (PubMed) https://pubmed.ncbi.nlm.nih.gov/40196638/

Funnelbeaker culture, Wikiwand, Updated 2 Jul 2025, https://www.wikiwand.com/en/articles/Funnelbeaker_culture

Corded Ware Culture – Origins and Development, Liquidsearch, https://www.liquisearch.com/corded_ware_culture/origins_and_development

[35] Out, Welmoed, The scale of human impact at the Hazendonk, the Netherlands, during the Late Neolithic, 153-165 in Harry Fokkens, Bryopny J. Coles, Annelou L. Van Gijn Jos P Kleijne, Hedwig H. Ponjee and Corijanne G. Slappendel, (eds) Between Foraging and Farming, Leiden: Leiden University 2008, https://www.researchgate.net/publication/236142152_The_scale_of_human_impact_at_the_Hazendonk_the_Netherlands_during_the_Late_Neolithic

[36] Altena E, Smeding R, van der Gaag KJ, Larmuseau MHD, Decorte R, Lao O, Kayser M, Kraaijenbrink T, de Knijff P. The Dutch Y-chromosomal landscape. Eur J Hum Genet. 2020 Mar;28(3):287-299. doi: 10.1038/s41431-019-0496-0. Epub 2019 Sep 5. Erratum in: Eur J Hum Genet. 2020 Mar;28(3):399. doi: 10.1038/s41431-019-0528-9. PMID: 31488894; PMCID: PMC7029002. (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC7029002/

Welmoed A. Out, Integrated archaeobotanical analysis: Human impact at the Dutch Neolithic wetland site the Hazendonk, Journal of Archaeological Science, Volume 37, Issue7, 2010, Pages 1521-1531, ISSN 0305-4403, https://doi.org/10.1016/j.jas.2010.01.012.
(https://www.sciencedirect.com/science/article/pii/S0305440310000178 )

Luc Amkreutz, Leo Verhart, Milco Wansleeben, Hazendonk layers over and over again, 139-151, in Harry Fokkens, Bryopny J. Coles, Annelou L. Van Gijn Jos P Kleijne, Hedwig H. Ponjee and Corijanne G. Slappendel, (eds) Between Foraging and Farming, Leiden: Leiden University 2008, https://www.academia.edu/7731906/Hazendonk_layers_over_and_over_again

Erik Drenth, Otto Brinkkemper and Roel C. G. M. Lauwerier, Amersfoort, Single Grave Culture Settlements in the Netherlands: the state of affairs anno 2006, in W.Dörfler, J. Müller, eds, Wirtshaft – Sied lung en im dritte n vorc hris tlic hen Ja hr-tauase nd Mitteleur opas und Süd skandinavi ens, Offa-Bücher 84 (Neum ünster 200 8) 149 -181, https://www.researchgate.net/publication/292249638_Umwelt-Wirtschaft-Siedlungen_im_dritten_vorchristlichen_Jahrtausend_Mitteleuropas_und_Sudskandinaviens

[37] Funnelbeaker Culture, Wikipedia, This page was last edited on 15 July 2025, https://en.wikipedia.org/wiki/Funnelbeaker_culture

Lipson M, Szécsényi-Nagy A, Mallick S, Pósa A, Stégmár B, Keerl V, Rohland N, Stewardson K, Ferry M, Michel M, Oppenheimer J, Broomandkhoshbacht N, Harney E, Nordenfelt S, Llamas B, Gusztáv Mende B, Köhler K, Oross K, Bondár M, Marton T, Osztás A, Jakucs J, Paluch T, Horváth F, Csengeri P, Koós J, Sebők K, Anders A, Raczky P, Regenye J, Barna JP, Fábián S, Serlegi G, Toldi Z, Gyöngyvér Nagy E, Dani J, Molnár E, Pálfi G, Márk L, Melegh B, Bánfai Z, Domboróczki L, Fernández-Eraso J, Antonio Mujika-Alustiza J, Alonso Fernández C, Jiménez Echevarría J, Bollongino R, Orschiedt J, Schierhold K, Meller H, Cooper A, Burger J, Bánffy E, Alt KW, Lalueza-Fox C, Haak W, Reich D. Parallel palaeogenomic transects reveal complex genetic history of early European farmers. Nature. 2017 Nov 16;551(7680):368-372. doi: 10.1038/nature24476. Epub 2017 Nov 8. PMID: 29144465; PMCID: PMC5973800. (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC5973800/

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Erik Drenth, Otto Brinkkemper and Roel C. G. M. Lauwerier, Amersfoort, Single Grave Culture Settlements in the Netherlands: the state of affairs anno 2006, in W.Dörfler, J. Müller, eds, Wirtshaft – Sied lung en im dritte n vorc hris tlic hen Ja hr-tauase nd Mitteleur opas und Süd skandinavi ens, Offa-Bücher 84 (Neum ünster 200 8) 149 -181, https://www.researchgate.net/publication/292249638_Umwelt-Wirtschaft-Siedlungen_im_dritten_vorchristlichen_Jahrtausend_Mitteleuropas_und_Sudskandinaviens

[38] van den Dikkenberg, Lasse, Vlaardingen culture or Stein group? From object biographies to communities of practice, in: Verbaas, A., Langejans, G., Little, A. and Chan, B. (eds). Artefact Biographies from Mesolithic and Neolithic Europe and Beyond. Papers in honour of Professor Annelou van Gijn. Analecta Praehistorica Leidensia 52. Leiden: Sidestone Press, pp. 63-72, 6 Jun 2024,  DOI: 10.59641/pp090sb https://www.researchgate.net/publication/381409197_Vlaardingen_culture_or_Stein_group_From_object_biographies_to_communities_of_practice_in_Verbaas_A_Langejans_G_Little_A_and_Chan_B_eds_Artefact_Biographies_from_Mesolithic_and_Neolithic_Europe_and_Bey

Van Den Dikkenberg, Lasse and Alex Brandsen, Using Text Mining to Search for Neolithic Vlaardingen Culture Sites in the Rhine-Meuse-Scheldt Delta, Journal of Computer Applications in Archaeology, 8(1): 110–124. DOI: https://doi.org/10.5334/jcaa.205 , https://journal.caa-international.org/articles/205/files/67e14353e173f.pdf

Erik Drenth, Otto Brinkkemper and Roel C. G. M. Lauwerier, Amersfoort, Single Grave Culture Settlements in the Netherlands: the state of affairs anno 2006, in W.Dörfler, J. Müller, eds, Wirtshaft – Sied lung en im dritte n vorc hris tlic hen Ja hr-tauase nd Mitteleur opas und Süd skandinavi ens, Offa-Bücher 84 (Neum ünster 200 8) 149 -181, https://www.researchgate.net/publication/292249638_Umwelt-Wirtschaft-Siedlungen_im_dritten_vorchristlichen_Jahrtausend_Mitteleuropas_und_Sudskandinaviens

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Amkreutz, Luc, Funerary Practices on the Fringe, 21-33, in  Daniela Hofman, Vicki Cummongs, Mathias Bjørnevad-Ahlqvist and Rune Iversen, eds, The Early Neolithic of Northern Europe, Leiden: Sidestone Press Academics, 2025   https://www.sidestone.com/books/the-early-neolithic-of-northern-europe

Brinkkemper, Otto, Erik Drenth and Jørn Zeiler, An outline of the subsistence of the Vlaardingen culture from the Netherlands, Revue archéologique de Picardie, Année 2011 NS 28 pp. 207-220, https://www.persee.fr/doc/pica_1272-6117_2011_hos_28_1_3331

Limburg (Netherlands), Wikipedia, This page was last edited on 7 June 2025, https://en.wikipedia.org/wiki/Limburg_(Netherlands)

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[40] Cassen, Serge (2019). “Real and ideal European maritime transfers along the Atlantic coast during the Neolithic” (https://www.academia.edu/54283882). Documenta Praehistorica. 46. doi:10.4312/dp.46 (https://doi.org/10.4312%2Fdp.46)

Michelsberg culture (4400 to 3500bc) is characterized by undecorated pointy based tulip beakers, https://neolithicarch.com/michelsberg-culture-4400-to-3500bc-is-characterized-by-undecorated-pointy-based-tulip-beakers/

Darvill, Timothy, ed. (2009). “Michelsberg Culture”. The Concise Oxford Dictionary of Archaeology (3 ed.). Oxford University Press. ISBN9780191727139.

Michelberg Culture, Wikipedia, This page was last edited on 24 December 2024, https://en.wikipedia.org/wiki/Michelsberg_culture

Funnelbeaker culture, Wikiwand, Updated 2 Jul 2025, https://www.wikiwand.com/en/articles/Funnelbeaker_culture

Folkens, Harry, Background to Dutch Beakers. A Critical Review of the Dutch Model, Pages 9 – 36 in Harry Fokkens and Franco Nicolis, eds, Background to Beakers, van Woerdekom: Sidestone Press, 2012, 
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Funnelbeaker culture, Wikiwand, Updated 2 Jul 2025, https://www.wikiwand.com/en/articles/Funnelbeaker_culture

Folkens, Harry, Background to Dutch Beakers. A Critical Review of the Dutch Model, Pages 9 – 36 in Harry Fokkens and Franco Nicolis, eds, Background to Beakers, van Woerdekom: Sidestone Press, 2012, 
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[42] Tanja Zerl, Manfred Rösch, and Elena Marinova, The Late Neolithic Michelsberg culture – just ramparts and ditches? A supraregional comparison of agricultural and environmental data, Prähistorische Zeitschrift 89(1), 2014, https://doi.org/10.1515/PZ-2014-0006

Crombé, P., Aluwé, K., Boudin, M. et al. New evidence on the earliest domesticated animals and possible small-scale husbandry in Atlantic NW Europe. Sci Rep 10, 20083 (2020). https://doi.org/10.1038/s41598-020-77002-4

[43] Michelberg Culture, Wikipedia, This page was last edited on 24 December 2024, https://en.wikipedia.org/wiki/Michelsberg_culture

[44] Tanja Zerl, Manfred Rösch, and Elena Marinova, The Late Neolithic Michelsberg culture – just ramparts and ditches? A supraregional comparison of agricultural and environmental data, Prähistorische Zeitschrift 89(1), 2014, https://doi.org/10.1515/PZ-2014-0006

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Teetaert D, Baeyens N, Perdaen Y, et al. A well-preserved Michelsberg Culture domed oven from Kortrijk, Belgium. Antiquity. 2019;93(368):342-358. doi:10.15184/aqy.2018.181 https://www.cambridge.org/core/journals/antiquity/article/abs/wellpreserved-michelsberg-culture-domed-oven-from-kortrijk-belgium/36403BAC5E60E0CE807C840BB7F6BF43

[45] Vlaardingen culture, Wikipedia, This page was last edited on 21 June 2025, https://en.wikipedia.org/wiki/Vlaardingen_culture

Van Den Dikkenberg, Lasse and Alex Brandsen, Using Text Mining to Search for Neolithic Vlaardingen Culture Sites in the Rhine-Meuse-Scheldt Delta, Journal of Computer Applications in Archaeology, 8(1): 110–124. DOI: https://doi.org/10.5334/jcaa.205 , https://journal.caa-international.org/articles/205/files/67e14353e173f.pdf

[46] Folkens, Harry, Background to Dutch Beakers. A Critical Review of the Dutch Model, Figure 5 The Cultural Landscape of the Netherlands c. 3400-2850 cal BC, Pages 9 – 36 in
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[47] Menne, Julia and Mirco Brunner, Transition from Swifterbant to Funnelbeaker: A Bayesian Chronological Model, Open Archeology, Oct 2021 7(1): 1235-1243 DOI:10.1515/opar-2020-0191
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[48] Funnelbeaker culture, Wikipedia, This page was last edited on 2 July 2025, https://en.wikipedia.org/wiki/Funnelbeaker_culture

Corded Ware culture, Wikipedia, This page was last edited on 23 June 2025, https://en.wikipedia.org/wiki/Corded_Ware_culture

Olalde I, et al , Long-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups. bioRxiv [Preprint]. 2025 Mar 25:2025.03.24.644985. doi: 10.1101/2025.03.24.644985. PMID: 40196638; PMCID: PMC11974744 (PubMed) https://pubmed.ncbi.nlm.nih.gov/40196638/

Funnelbeaker culture, Wikiwand, Updated 2 Jul 2025, https://www.wikiwand.com/en/articles/Funnelbeaker_culture

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[49] Sánchez-Quinto F, Malmström H, Fraser M, Girdland-Flink L, Svensson EM, Simões LG, George R, Hollfelder N, Burenhult G, Noble G, Britton K, Talamo S, Curtis N, Brzobohata H, Sumberova R, Götherström A, Storå J, Jakobsson M. Megalithic tombs in western and northern Neolithic Europe were linked to a kindred society. Proc Natl Acad Sci U S A. 2019 May 7;116(19):9469-9474. doi: 10.1073/pnas.1818037116. Epub 2019 Apr 15. PMID: 30988179; PMCID: PMC6511028. (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC6511028/

[50] Chyleński, M., Makarowicz, P., Juras, A. et al. Patrilocality and hunter-gatherer-related ancestry of populations in East-Central Europe during the Middle Bronze Age. Nat Commun 14, 4395 (2023). https://doi.org/10.1038/s41467-023-40072-9 https://www.nature.com/articles/s41467-023-40072-9

[51] Dolmens are ancient megalithic structures typically consisting of two or more large upright stones supporting a massive horizontal stone slab, often interpreted as tombs.

Dolmens, Wikipedia, This page was last edited on 19 July 2025, https://en.wikipedia.org/wiki/Dolmen

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[53] Funnelbeaker culture, Wikiwand, Updated 2 Jul 2025, https://www.wikiwand.com/en/articles/Funnelbeaker_culture

Funnelbeaker culture, Wikipedia, This page was last edited on 2 July 2025, https://en.wikipedia.org/wiki/Funnelbeaker_culture

Sánchez-Quinto F, Malmström H, Fraser M, Girdland-Flink L, Svensson EM, Simões LG, George R, Hollfelder N, Burenhult G, Noble G, Britton K, Talamo S, Curtis N, Brzobohata H, Sumberova R, Götherström A, Storå J, Jakobsson M. Megalithic tombs in western and northern Neolithic Europe were linked to a kindred society. Proc Natl Acad Sci U S A. 2019 May 7;116(19):9469-9474. doi: 10.1073/pnas.1818037116. Epub 2019 Apr 15. PMID: 30988179; PMCID: PMC6511028. (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC6511028/

[54] Funnelbeaker culture, Wikiwand, Updated 2 Jul 2025, https://www.wikiwand.com/en/articles/Funnelbeaker_culture

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[55] Kroon, E. J., Serial learners: interactions between Funnel Beaker West and Corded Ware communities in the Netherlands during the third millennium BCE from the perspective of ceramic technology. Sidestone Press, Leiden. 18 Sep 2024 Retrieved from https://hdl.handle.net/1887/4092578 ; https://scholarlypublications.universiteitleiden.nl/access/item:4092595/view

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