Looking at the Griff(is)(es)(ith) Y-DNA Phylogenetic Gap Associated with the Meuse and Rhine River Watershed from the Bronze Age Onward – Part Four

This fourth part of the story focuses on possible influences during and after the bronze age, up to approximately 650 CE, the estimated birth date of the most recent common ancestor that is asociated with haplogroup G-7648 at the end of the phylogenetic gap. Some of these influences are:

• The enduring effects of the Bronze Age Bottleneck;
• The enduring impact of the Corded Ware and Bell Beaker migrations on limiting the proliferation of G haplogroup subclades in susequent generations; and
• Environmental impacts and changes in the delta landscape.

The 2,850 year Gap between G-FGC7516 and G-Z6748:  The most common recent ancestor associated with G-FGC7516 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 of undocumented YDNA ancestors represents about 95 generations.

The Enduring Impact of YDNA Demographic Patterns Restricting G2a Subclade Proliferation in the Muese Rhine Watershed Area

1. Unique Persistence and Admixture of Hunter-Gatherer and Farming Y-Haplogroups

The Meuse-Rhine watershed area exhibited a distinctive demographic pattern compared to much of Europe when early G2a haplogroup farmer groups initially migrated into the area. There was a significant, long-term persistence of local hunter-gatherer YDNA ancestry. For thousands of years, the Rhine-Meuse region (covering the western and central Netherlands, Belgium, and western Germany) maintained a population with very high hunter-gatherer ancestry, up to fifty percent, much longer than surrounding areas, due to limited integration of early farmer ancestry (e.g. G2a lineages). The unique persistence and admixture of hunter-gatherer and farming Y-haplogroups is an unique characteristic of the Meuse-Rhine area throughout the Neolithic and Chalcolithic periods. [1]

The Y-DNA haplogroups representing the local mix of European hunter-gatherers and early European farmers were primarily I2, C1, and G2a. These YDNA lineages reflect the blending of Mesolithic hunter-gatherer males and Neolithic agriculturalist males before the large influx of Steppe ancestry in the Bronze Age. The main I2 Y-DNA lineage was found among Western, Central, and Eastern European Mesolithic hunter-gatherers. They persisted widely through the Neolithic and into the Copper Age, especially in areas with continued high hunter-gatherer ancestry. Haplogroup C1 was much rarer haplogroup but was detected in some Western and Northern European Mesolithic and Early Neolithic remains, showing deep Paleolithic roots in Europe. [2]

The G2a Early European Farmer (EEF) male lineage, the lineage representing the Griff(is)(es)(ith) paternal line, was among the Anatolian/Levantine-derived early Neolithic farmers who spread agriculture into Europe through northern and sourthern Europe, especially in Linearbandkeramik (LBK), Cardial, and other early Neolithic cultures. [3] The F and J haplogroups, less common but also observed among some early farmer groups, were present particularly in southern and southeastern Europe. [4]

As mentioned in the video above, regions such as the Meuse-Rhine area, Central Europe, and France showed intermixed communities during the Neolithic and Chalcolithic periods, with both I2 (hunter-gatherer) and G2a/F (farmer) haplogroups detected among males. The presence of both I2 and G2a haplogroups in later Neolithic era sites indicates communities with significant admixture between local hunter-gatherers and arriving farming peoples.

In regions with high western hunter-gatherer and early European farmer admixture, especially before the steppe-related R1b spread, Y-DNA lineages I2 and G2a were typical representatives of the local male genetic landscape. As reflected in table one, ancient DNA (aDNA) from the Neolithic and Chalcolithic Meuse-Rhine region shows that males predominantly belonged to haplogroups I2a, R1b-V88, and C1a—all linked to European hunter-gatherers [5]

Table One: Haplogroup Presence in the Mesue Rhine Watershed Area

Period/Population	Y-DNA Haplogroups	Description
Mesolithic	I2a, C1a, R1b-V88	Western Hunter-gatherer (WHG) dominant
Early Neolithic Mix	I2a, C1a, R1b-V88 + minor representation of G2a	40–50% WHG, 50–60% Early Eurpean Farmer (EEF) admixture
Post-Neolithic Beaker	R1b-L151	Steppe ancestry dominant, local mix wanes

The G2a haplogroup was present in the local Neolithic and Chalcolithic populations of the Meuse-Rhine area, but it was not as dominant there as in many other parts of Europe. While early European farmers across the continent are strongly associated with G2a, ancient DNA from the Rhine-Meuse region during the Neolithic instead shows a much higher persistence of hunter-gatherer lineages such as I2a and C1a, with G2a present but at relatively lower frequency. This may partly explain the dearth of discovered subclades in the Griff(is)(es)(ith) paternal migratory line of descent through this area of western Europe.

“The great winner during the Neolithic period was haplogroup I2a, which consistently shows up alongside G2a in most Neolithic sites tested to date (Starčevo, Körös, Lengyel, LBK, Cardium Pottery, Megalithic), and seem to increase in frequency over time and as one moves towards Northwest Europe.“

“(A)lthough I2a was just one of many Mesolithic hunter-gatherers’ lineages in Europe when agriculturists arrived, it is the only one that readily embraced the new lifestyle and managed to supersede the original farmers in number. I2a’s destiny was not only linked to its ability to chum with G2a, but we could say that G2a farmers catalysed I2a’s success. I2a people integrated G2a tribes, learned the new Neolithic techniques from them and became so good at them that over time the student overtook the master.“ [6]

The Rhine-Meuse region’s river-dominated landscapes significantly shaped the adoption and adaptation of Neolithic farming practices through ecological constraints, specialized subsistence strategies, and the blending of hunter-gatherer and farming cultural practices. The dynamic wetland/riverine environment (marshes, peat bogs, and seasonal floods) hindered full-scale Neolithic agriculture. Communities developed a mixed subsistence strategy combining:

• Limited crop cultivation on elevated river dunes/levees;
• Cattle husbandry optimized for wetland conditions (grazing on salt marshes, occasional winter foddering); and
• Persistent hunting, fishing, and foraging in resource-rich aquatic ecosystems.

This “extended broad spectrum” approach allowed populations to exploit the landscape without abandoning traditional Mesolithic practices. [7]

2. Impact of Bell Beaker Culture Migration and YDNA Replacement: A Local Admixture Event Resulting in New Populations with Dominant Steppe Ancestry

During the arrival of the Corded Ware complex, local individuals in this region adopted elements of the material culture but exhibited very little steppe ancestry, unlike Corded Ware sites elsewhere. Bell Beaker-associated populations in this region around 2500 BCE were formed by a mixture of Corded Ware-related migrants with steppe ancestry and the persistent local ‘Neolithic substrate’, with the local contribution modelled at 9 to 17 percent. Bell Beaker men in the Rhine-Meuse region mostly carried the R1b-L151 (especially P312) Y-chromosome, absent in earlier Neolithic populations but present among Central European Corded Ware groups, indicating a strong but not exclusive external influx. [8]

In much of Northwestern Europe, Bell Beaker and R1b haplogroup expansion involved almost total population and male lineage replacement. In contrast, as reflected in table two, the Rhine-Meuse area saw a transformative but partly local admixture event, resulting in new populations with dominant steppe ancestry (about 83–91 percent) but still a recognizable input from the enduring local hunter-gatherer- early European farmer influenced population. This “fusion zone” became a launching pad for further population expansions into regions such as Britain, where the Bell Beaker-associated transformation was even more complete. [9]

Table Two: Distinctive Demographic Impact in Muese Rhine Watershed Area

Region	Pre-Beaker Population	Beaker/Early Bronze Age Pattern	Estimated Ancestry Turnover	Local Genetic Input
Meuse-Rhine	Hunter-Gatherer ‘Rich Mix’ with Early European Farmers	Bell Beaker: R1b, steppe ancestry + local admixture	83–91% turnover	9–17%
Britain	Neolithic Farmers	Near-total replacement by R1b-rich, steppe Beaker	90–100% turnover	0–9%
Central Europe	Neolithic Farmers + WHG	R1b predominance, steppe ancestry via Corded Ware	Very high	Minimal

The Meuse-Rhine region stands out for its partially blended transformation during the Beaker phenomenon, marked by fusion rather than just replacement, with steppe-derived R1b Y-haplogroups and ancestry still predominating in the end. This admixture pattern was unusual for Europe. In most regions, Neolithic migration led to the overwhelming dominance of farmer-origin lineages (e.g. G2a), with much lower persistence of I2 or other hunter-gatherer Y-DNA. In the Meuse-Rhine, admixture continued for centuries, and the local hunter-gatherer Y-haplogroups persisted at high levels until the arrival of Bell Beaker people with predominantly steppe and R1b-L151 ancestry.

3. The Enduring Impact of Post-Bronze Age Influences

The decline of G2a subclades in the Netherlands was not a single event but a cumulative process over generations. It began with the Bronze Age migrations that replaced many Neolithic paternal lineages, and was further influenced by demographic, environmental, and social changes in subsequent time periods.

Pre-existing populations, including those with high frequencies of G2a, may have faced bottlenecks, diseases, or environmental changes that reduced their numbers, making them more susceptible to genetic replacement. For example, archaeological evidence from the Netherlands points to periods of population decline and settlement abandonment during the Roman Empire’s collapse, which further complicated and reshaped the genetic landscape. [10]

Genetic drift, the random fluctuation of gene variant frequencies in a population, would have also contributed to the decline of G2a. In small, isolated communities, the loss of certain lineages due to chance events or the failure of a male line to reproduce can have a significant impact. Shifts in reproductive dynamics, including new social structures and marriage patterns brought by the migrating populations, may have further disadvantaged the older G2a paternal lines. [12]

Subsequent migrations during the Iron Age and the early Middle Ages—such as the arrival of Germanic tribes (including Angles, Saxons, and Franks)—continued to shape the genetic makeup of the Low Countries in the Muese Rhine watershed area. This further diluted or replaced the genetic signatures of earlier groups, cementing the marginalization of older lineages like G2a. [13]

A recent study by Eveline Altena and associates provides an in-depth look at paternal genetic continuity in the Netherlands across a span of 1,300 years. The key findings indicate remarkable stability of male lineages. The team analyzed the Y-chromosomes of 348 men from 13 Dutch locations dated 500 CE–1850 CE, alongside modern YDNA data, to trace paternal ancestry across millennia, see illustration one. [14]

Illustration One: Observed Y-Haplogroup Frequencies in Three Historical Periods in the Middle Ages in the Netherlands

The study found that the male population of the Netherlands showed limited change from the Early Middle Ages to modern times. Based on the study’s findings, fluctuations in haplogroup frequencies mostly resulted from genetic drift rather than large-scale migration or population replacement.

The study also supports the impression that the medieval Netherlands practiced patrilocality—men stayed put in localized areas and women moved for marriage. Regional male lineages were reinforced and preserved by limited male mobility, while mitochondrial (female) variation was more diffuse.

“The population substructure and gradients for many of the individual YHGs (Y haplogroups) we found in our study are in strong contrast with the apparent lack of genetic-geographic patterns for mtDNA data . . . . This could be an indication of different demographic histories for women and men. One could think, for example, of the patrilocal residence system, which is typical for farming societies, such as the Dutch. In these societies sons stay with their family and daughters move to the residence of their husbands. Also, genetic drift may have acted differently on mt-DNA than on Y-chromosomes.” [15]

Regional male lineages were reinforced and preserved by limited male mobility, while mitochondrial (female) variation was more diffuse. Over centuries, Dutch male lineages were shaped less by mass migration than by chance and social practice, setting them apart from regions with more turbulent demographic histories. [16]

In an earlier published article in 2019 and 2020, Altena and associates completed a geographic analysis of Y-chromosome haplogroup (YHG) distribution across the Netherlands. Using data from 2,085 males and integrating information from northern Belgium, the study found distinct geographic patterns in Y-chromosome distribution, with multiple Y-haplogroups showing significant clinal frequency gradients (i.e., gradual changes in frequency across regions). [17]

While previous research found limited or no mitochondrial DNA (maternal-line) spatial patterns or substructure, the pronounced Y-chromosomal substructure points to different population histories for men and women in the Netherlands. Contrasting male and female lineage patterns suggest sex-biased demographic histories, with male lineages undergoing more pronounced geographic differentiation, perhaps reflecting historical migration, social, or cultural practices.

Prediction surface maps were used to visualize the complex distributions of individual Y-haplogroups in detail, revealing non-random patterns throughout the country for almost every haplogroup examined. Y-chromosome diversity in the Netherlands shows a significant micro-geographic structure, with several haplogroups (e.g., R1b variants) displaying regional gradients.

Illustration Two: Prediction Surface Maps of the Four Most Frequent Y Haplogroups in the Dutch Dataset

Y-haplogroups “G-M201, J2-M172, R1b-M269, and R1b-S116 increase from north to south, R1b-M405 Total and R1b-L48 increase from south to north, I-M170 increases from southwest to northeast and R1b-S116 Total, R1b-U152 and R1b-M529 increase from northeast to southwest. . . . From all the YHGs for which prediction surface maps were created, only YHG R1b-M529 is more or less evenly distributed over the Netherlands. All other YHGs show more distinct patterns of distribution.” [18]

Altena and associate researchers point out that the haplogrup G-M201 which the Griff(is)(es)(ith) paternal line is part of “is relatively rare in Europe, with average proportions below 5% in northwestern Europe, which is consistent with our findings. Because proportions are low throughout the most of Europe, there is no clear gradient, but overall it increases from northwest to southeast (in the Netherlands and Belgium) . . . .“. [19]

Today’s clear-cut geographic patterns in Dutch Y-DNA formed late, likely as the result of more recent events and not from deep medieval roots. [20]

The Use of Archaeological Time Periods Associated with this Phylogenetic Gap

The loose reference at the begining of this story to a time period “during and after the bronze up to 650 CE” can be referenced by and viewed through the following archaeological time periods in table three. The approximate dates for the three Bronze Age periods, the two periods of the Iron Age, the Roman era and the Merovian perod in the Meuse-Rhine river watershed are based on archaeological and historical studies. These approximate time ranges reflect scholarly consensus for distinctive time periods for the Meuse-Rhine watershed, though there may be slight regional variation within the area.

Utilizing archaeological time periods offers the advantage of organizing human history into discernible phases. Archaeological time periods (like the Stone, Bronze, and Iron Ages) help organize the development of human societies in a clear chronological framework, making it easier to compare changes across time and regions. These periods enable archaeologists to interpret artifacts and features within a broader cultural and technological context, offering insight into the evolution of societies, technologies, and economies. Standardized periods facilitate the communication of archaeological findings to both academic communities and the public, enhancing understanding of complex historical developments. [21]

However, there are obvious limitations associated with the use of archaeological time periods. Broad periods like “Stone Age” or “Iron Age” can mask local and regional diversity, as technological advancements and cultural practices often occurred at different times in different places, leading to overlaps that the periods cannot accurately reflect. Assigning boundaries to these periods involves subjective decisions based on material culture, which can introduce bias and debate among scholars. [22]

Table Three: Archaeological Periods in the Meuse Rhine Watershed Area During the Phylogentic Gap for the Griff(is)(es)(ith YDNA Lineage

Archaeological Period	Approximate Dates	Archeaological Description
Early Bronze Age	2200–1600 BCE	At the beginning of the Bronze Age, the Meuse-Rhine area was at a cultural crossroads and was influenced by broader European traditions. The Beaker culture dominated much of Western and Central Europe and is considered a transitional culture between the late Neolithic and the early Bronze Age. While primarily present further east, the Corded Ware culture also influenced the early Bronze Age in parts of the region, with archaeologists sometimes tracing burial rituals back to these roots.  Communities were settled along elevated stream ridges and river banks, engaging in mixed farming and livestock herding, reflecting the broader transition to more sedentary life seen kin other parts of north and central Europe. There was the persistence of distinctive local cultures with high levels of hunter-gatherer ancestry, in contrast to the rest of Europe where farming ancestry became dominant earlier. The area’s wetlands and rivers promoted more isolated populations with limited early integration of outside groups. [23]
Middle Bronze Age	1600–1300 BCE	The region experienced a significant increase in settlements, with numerous house sites discovered in the river delta, facilitated by technological advances such as improved bronze tools and animal husbandry. Elp culture (northern sector): Located in the northern and eastern Netherlands, this culture existed from approximately 1800 to 800 BCE. Hilversum culture (southern sector): Centered in the southern Netherlands and northern Belgium, this culture has been linked to the Wessex culture of southern England. Long-distance contacts and cultural exchange increased, reflected in the archaeological record by the emergence of ‘pan-European’ material culture, but settlements declined toward the end of this period hints at possible floods or ecological stresses. Large-scale deforestation for agriculture is evident, paralleling the broader trend of expanding farmland and increased social complexity. [24]
Late Bronze Age	1300–800 BCE	The Urnfield culture became the dominant cultural tradition across Central Europe and extended into the Meuse-Rhine area during the Late Bronze Age. A local sub-group known as the Lower Rhine Group developed within the Urnfield tradition. The Urnfield period saw an increase in social stratification and widespread trade networks. It is also characterized by the appearance of fortified hilltop settlements and advanced bronze metalworking. A sharp decline in settlements is noted in the region, partly attributed to climate change, flooding, and changes in river dynamics, but also possibly to changing social structures or shifts toward more mobile ways of life. Evidence points to sustained but reduced habitation, with human activity focused on certain elevated or defendable sites. This aligns with the general Bronze Age trend toward urbanization and sociopolitical stratification, but environmental instability made large, permanent settlements less sustainable. There was significant contact with neighboring areas, visible in material culture and genetics, corresponding with the general increase in social complexity and long-distance exchange in the wider Bronze Age world. [25]
Iron Age [26]	Early Iron Age (Hallstatt period): 800-500 BCE Late Iron Age (La Tène period): 450 -50 BCE	The Meuse-Rhine area served as a transitional zone between the Celtic south (La Tène) and the Germanic north (Hallstatt), resulting in a unique blend of traditions. The area saw continuity from prehistoric populations with new influences entering gradually, rather than abrupt cultural shifts. Settlements in the Meuse-Rhine Iron Age area were typically small and lacked the major fortified oppida found further south in La Tène regions. Archaeological surveys note a decline or transformation in settlement patterns in the early Iron Age, with fewer large settlements and more dispersed smaller sites, contrasting with the denser occupation of the Late Bronze Age. [27] The region experienced significant environmental changes such as increased flooding and sedimentation in the river valleys, likely due in part to enhanced human land use such as farming and deforestation. Archaeological evidence shows locally produced material goods, e.g. pottery, basic iron tools, and simple burial rituals rather than elaborate grave goods. The adoption of La Tène cultural elements—occurred only to a limited degree, reflecting modest trade and cultural interaction with Celtic areas.
Roman Era [28]	55 BCE–459/461 CE	The Roman Era in the Meuse-Rhine Watershed was defined by advances in water management, the creation of complex settlements along the Roman frontier, multicultural dynamics, and a blend of indigenous and Romanized landscape use, leaving a distinctive archaeological legacy.
Late Antiquity/Post-Roman Transition	460–500 CE	Collapse of Roman state authority, significant depopulation, urban and rural structures deteriorated, the Roman limes (frontier) ceased to function [29]
Early Merovingian Period	500–650 CE	Region became part of Frankish kingdom, under Merovingian rule. Settlement patterns shifted, with new rural settlements developing—sometimes continuing from or near former Roman sites—and local communities reorganizing around new Frankish leadership structures. The area retained the Roman “pagi” (district) administrative structure, following both old Roman and Germanic tribal traditions in leadership and justice, with leaders initially selected by local warriors. The chaotic social context, continually punctuated by small-scale warfare, fostered the rise of a local warrior elite—the precursors to later medieval nobility—rewarded with land for military service. [30]

While archaeological time periods help bring order and clarity to the study of human prehistory and history, they are best used as flexible frameworks rather than rigid chronological boundaries, always considering the limitations and regional variations they entail. Regional chronologies—such as division into Early, Middle, and Late Bronze Age—are not uniform and may overlap or differ in exact dating due to differing archaeological traditions and the tempo of cultural change and adoption of technicalogcal change as reflected in archaeological artifacts.

The limitations of utilizing ‘time periods’ can be specifically demonstrated when looking at the ‘bronze age’ in the Meuse-Rhine watershd area. The Bronze Age in Europe shows significant regional variation in both its starting and ending dates, as well as in developmental phases. The earliest Bronze Age developments in Europe occurred in the Aegean and southeast Europe, spreading gradually northwest and north. Central and northern regions (e.g. Scandinavia and the British Isles) entered the Bronze Age later due to environmental, technological, and cultural factors.

Archaeologist Harry Fokkens has called for a reassessment of the criteria used to define archaeological periods, advocating for a more multi-faceted approach that combines absolute carbon dating with meaningful archaeological markers, such as changes in settlement organization, technology, and social structure. He urges, as summarized in table two, archaeologists to move beyond simply updating dates and to critically reflect on the cultural significance of period boundaries (see table four). [31]

Table Four: Fokkens’ Recommendations for Sustantiation of Archeological Periods

Aspect	Current Practice	Fokkens’ Proposed Revision
Primary Evidence	Burial data and bronzes	Inclusion of Settlement patterns, houses, material culture
Theoretical Basis	Migration/culture-change models	Localized cultural continuity analysis
Chronological Tools	Radiocarbon dating	Combined: carbon dating & cultural/economic shifts
Transparency	Assumptions rarely scrutinized	Explicit criteria and peer-reviewed frameworks

Fokkens argues that the division of the Bronze Age into different cultures and phases was predominantly based on burial data, the nature and use of pottery, and migration theories and culture-change models that often relied on weak evidence for social, religious, or spiritual changes, which, while detailed, lacked scientific credibility. Other cultural phenomena like settlement patterns and house types were underutilized or inconsistently considered. He argues that the bronze age in the Netherlands existed roughly between 1800 – 800 BCE.

” … (W)e should move the line dividing the Neolithic and Bronze Age towards 1800 BC. This is the absolute end of the Beaker traditions and the start of entirely new ones, maybe not so much in a technological sense but in social, economical and ritual aspects of culture. I therefore propose to indicate the period between 2900 and 1800 BC as the Late Neolithic, the period of the Beaker Cultures. Late Neolithic A is the period of the Single Grave Culture including AOO Beakers (2900-2500 BC), Late Neolithic B the period of the Bell Beaker Culture (2500- 2000), Late Neolithic C the period of the Barbed Wire Beaker Culture (2000-1800). …

“The Early Bronze Age, as I see it, is marked by the development of several traditions that differ from the Late Neolithic practices sufficiently to suggest that a fundamental change in several dimensions of culture occurred simultaneously. Housing traditions changed and possibly associated economic traditions, burial traditions, deposition practices and pottery traditions. At the same time new traditions start which continue for the next 700 years.” [32]

While Fokkens does not propose a specific alternative chronological framework with fixed phases and dates for the Dutch Bronze Age, the chronological boundaries of the Bronze Age in the Netherlands are generally considered to span from around 2000 BCE to 800 BCE based on research studies and archaeological periodization. While some sources mention minor earlier traces or slight local variation, most scholarly consensus places the start at about 2000 BCE, marked by the first appearance of bronze objects and the Barbed-Wire Beaker culture, and the end at around 800 BCE, when the transition to the Iron Age occurs.

Martin Furholt, makes a similar agument when discussing the European Neolithic Period. He highlights how recent advances in archaeogenetics—the analysis of ancient DNA—have revolutionized the understanding of mobility and social dynamics in Neolithic Europe. He also critically examines the interpretations and intellectual frameworks that have emerged from this new data. [33]

Furholt argues that the enthusiasm for genetic data may lead to simplistic, macro-level migration narratives (e.g., mass migrations or population replacements), reminiscent of earlier, now outdated models from the early 20th century that treated archaeological cultures as homogenous, biologically-defined groups. He stresses the importance of integrating genetic findings with archaeological and anthropological theory, warning against equating genetic patterns directly with social identity or assuming that genetic groups correspond neatly to archaeological cultures.

Furholt recommends moving beyond the idea of migration as a simple mass movement and instead encourages nuanced, case-by-case studies that combine diverse lines of evidence, such as material culture, burial practices, and genetic data. Furholt calls for better interdisciplinary collaboration between archaeologists and geneticists so that social, cultural, and biological data can together produce more sophisticated and accurate models of Neolithic social change. He calls for better interdisciplinary collaboration between archaeologists and geneticists so that social, cultural, and biological data can together produce more sophisticated and accurate models of Neolithic social change.

Bronze Age Cultures in the Muese-Rhine Watershed

The Bell Beaker phenomenon was never a monolithic culture but rather a complex network of related groups with substantial regional variation. As Bronze Age societies developed, these regional groups either merged with or were absorbed by neighboring cultures, leading to the fragmentation and eventual disappearance of distinct Bell Beaker identities. Key cultures marking the start and end of the European Bronze Age differ by region but show several clear patterns. [34]

The exact cause of the decline of the Bell Beaker culture is a subject of ongoing debate among archaeologists and historians, and there are various theories proposed. It is generally understood that the Bell Beaker culture, which flourished in Europe from around 2800 to 1600 BCE, eventually faded and was succeeded by new cultural norms and the onset of the Bronze Age. [35]

The Bell Beaker culture did not vanish abruptly. It was succeeded by emerging Bronze Age cultures such as the Únětice culture in Central Europe and various regional Bronze Age societies elsewhere. This transition often involved the blending of Bell Beaker traditions with those of incoming or neighboring groups, leading to new cultural identities and practices.

The major cultural groups succeeding the Bell Beakers in the Meuse-Rhine watershed were the Hilversum Culture during the Early Bronze Age, followed by the Urnfield Culture in the Middle to Late Bronze Age, leading into the various Early Iron Age societies. As the Urnfield Culture transitions to the Early Iron Age, archaeological evidence suggests further shifts, possibly involving early Hallstatt influences and precursor groups to the Celts (see table five). It is likely that the roughly 95 undocumented generations of the Griff(is)(es)(ith) paternal line were part of many of the cultures listed in table five.

Table Five: Major Cultural Groups in the Meuse Rhine Watershed Area by Archaeological Period

Culture	Period (BCE)	Archaeological Period and Location
Corded Ware	~ 2900–2400	Late Neolithic/Early Bronze Age
Bell Beaker	~ 2500–2100	Early Bronze Age; mix of local/steppe ancestry
Barbed Wire Beaker	~ 2000–1800	Early Bronze Age – Late Bell Beaker variant in region
Hilversum	~ 1870–1050	Middle Bronze Age, south Netherlands/Belgium
Elp	~ 1800–800	Middle to Late Bronze Age, north/east Netherlands
Tumulus	~ 1600–1200	Middle Bronze Age, tumulus (barrow) burials
Lower Rhine Urnfield	~ 1300–750	Late Bronze Age, cremation fields
Hallstatt	~ 800–450	Early Iron Age (C/D proper); outgrowth of Urnfield, proto-Celtic
La Tène, Germani cisrhenani	~ 450 – 50	Late Iron Age
Batavi, Cananefates, Tungri, Romanized locals	~ 50 BCE – 400 CE	Roman Era
Frankish (merovingian), Saxons, Frisians	~ 400 – 800 CE	Early Medieval

These cultural transformations set the stage for later groups in the Iron age such as the Nordwestblock, theorized as a distinct cultural area between Celtic and Germanic zones in the Lower Rhine and Meuse area. [36]

“The Bell Beaker cultures (2700–2100) locally developed into the Bronze Age Barbed-Wire Beaker culture (2100–1800). In the second millennium BC, the region was the boundary between the Atlantic and Nordic horizons and was split into a northern and a southern region, roughly divided by the course of the Rhine.“ [37]

The Rhine River would continue to have an ecological impact on competing cultural and social groups and have an effect on the YDNA composition through susequent archaeological time periods. It would also have an impact on the absence of documented YDNA subclades associated with the migratory path of the Griff(is)(es)(ith) paternal line.

Parting Comments

The contested Meuse and Rhine River watershed area shaped and restricted the growth of the Y-chromosomal DNA phylogenetic tree of the Griff(is)(es)(ith) paternal line through dynamic interactions between ecological barriers and other social groups. Hunter gatherer groups, male-biased migrations, demographic impacts on subclade proliferation and geopolitical conflicts in each of the archaeological times periods.

The region’s wetlands and riverine landscapes resisted early Neolithic farming, preserving I2a and C1a Y-haplogroups linked to Mesolithic hunter-gatherers and limiting subclade proliferation of G2a haplogroups. Genetic continuity persisted until about 2500 BCE, , longer than in continental Europe, due to limited integration of Anatolian farmer females into local communities. This ecological resistance created a “genetic refugium” for older paternal lineages.

Despite adopting Corded Ware cultural practices, lowland Rhine-Meuse populations retained minimal steppe ancestry. However, Corded Ware associated R1a-M417 Y-haplogroups were introduced through male-dominated migrations, while retaining about 50 percent of local Neolithic ancestry. Limited female gene flow from steppe populations preserved high hunter-gatherer autosomal ancestry.

The formation of Bell Beaker groups through the mixing of local Rhine-Meuse populations (about 9–17%) and Corded Ware migrants led to the near complete Y lineage replacement of G2a YDNA groups. Bell Beaker associated males carried R1b-L151 subtypes (e.g., P312, U106), replacing earlier Neolithic lineages such as the Griff(is)(es)(ith) YDNA line. Steppe-ancestry males again mixed with local females.

These patterns demonstrate how the Rhine-Meuse watershed’s contested history created a stratified YDNA phylogeny of dominant and minorty subclades, with clines and subclades reflecting a millennia of asymmetric gene flow, male-driven expansions, and ecological resilience.

An highly contested geographical region like the Meuse-Rhine watershed, coupled with its ecological influences, profoundly shaped phylogenetic YDNA trees over time, creating a genetic landscape marked by strong lineage turnover and historical layering. The unique combination of ecological complexity and frequent territorial disputes in this area led to several characteristic outcomes:

Persistent Ancient Minority Lineages Through Isolation

Swampy lowlands, river deltas, and ecological bottlenecks preserved ancient hunter-gatherer Y-haplogroups (such as I2a and C1a) for thousands of years longer than in neighboring regions. These ‘refugia effect zones’ resisted incoming male lineages from early farming and steppe cultures, leading to deep-rooted branches on local YDNA trees that survived alongside more recent YDNA subclades.

Abrupt Lineage Replacements via Male-Dominated Migrations

Periods of intense population turnover—such as the Corded Ware and Bell Beaker expansions—produced abrupt, near-complete Y-chromosome replacements marked by new dominant haplogroups like R1b-L151 and R1a-M417 in the archaeological record. These replacements rarely erased all earlier diversity, but pruned YDNA trees of specific Neolithic branches while fostering ‘star-like phylogenies’ indicative of founder effects and rapid expansions. [38]

Genetic Subclades Shaped by Political Borders

Long-standing borders acted as semi-permeable filters that constrained male gene flow. These boundaries, such as the Rhine and Meuse Rivers, imprint phylogenetic trees with localized subclades and long phylogentic trees with few branches, revealing the enduring effect of contested territories.

Frequent social and political fragmentation after various groups changed or collapsed magnified genetic drift and lineage bottlenecks in isolated settlements. This created micro-regional YDNA substructures, amplifyed minor lineages through stochastic processes while reducing overall haplogroup diversity in places with population contractions.

Accumulation of Genetic Layers in the ‘Long Duration’ of YDNA Genetic Time

As social groups vied for control over time, the region’s YDNA composition mirrored the area’s archaeological and historical complexity: every territorial upheaval left its signature as a branch or cluster on the YDNA phylogeny that may persist or be overwritten by subsequent events.

In combination, these impacts mean highly contested regions like the Meuse-Rhine watershed display phylogenetic YDNA trees marked by both ancient depth and recent star-phylogeny expansions, strong subregional differentiation, and evidence of recurrent lineage pruning and replacement driven by both ecological and sociopolitical forces.

The final part of this story will discuss the major cultural groups associated with the migratory path for the Griff(is)(es)(ith) paternal line succeeding the Bell Beakers through the Bronze Age, Iron Age, the Roman Era and the early medieval age.

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] 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, Theoten Anscher , Michel Toussaint, Kim Callan, Olivia Cheronet, Trudi Frost, Lora Iliev, Matthew Mah, Adam Micco, Jonas Oppenheimer, IrisPatterson, Lijun Qiu, Gregory Soos, J. Noah Workman, Ceiridwen J. Edwards, Losif 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, https://www.biorxiv.org/content/10.1101/2025.03.24.644985v1.full; also found at: https://pubmed.ncbi.nlm.nih.gov/40196638

[2] Szécsényi-Nagy Anna, Guido Brandt, Wolfgang Haak, Victoria Keerl, János Jakucs , Sabine Möller-Rieker, Kitti Köhler, Balász Gusztáv Mende, Krisztián Oross , Tibor Marton, Anett Osztás, Viktória Kiss, Marc Fecher, Gyögy Pálfi, Erika Molnár, Katalin Sebők, András Czene, Tibor Paluch, Mario Šlaus, Mario Novak, Nives Pećina-Šlaus, Brigitta Ősz, VandaVoicsek, Krisztina Somogyi, Gábor Tóth, Bernd Kromer, Eszter Bánffy, Kurt W. Alt. Tracing the genetic origin of Europe’s first farmers reveals insights into their social organization. Proc Biol Sci. 2015 Apr 22;282(1805):20150339. doi: 10.1098/rspb.2015.0339. PMID: 25808890; PMCID: PMC4389623 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC4389623/

António Faustino Carvalho, Eva Fernández-Domínguez, Eduardo Arroyo-Pardo, Catherine Robinson, João Luís Cardoso, João Zilhão, Mário Varela Gomes, Hunter-gatherer genetic persistence at the onset of megalithism in western Iberia: New mitochondrial evidence from Mesolithic and Neolithic necropolises in central-southern Portugal, Quaternary International, Volumes 677–678, 2023, Pages 111-120, ISSN 1040-6182,
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Primorac D, Šarac J, Havaš Auguštin D, Novokmet N, Bego T, Pinhasi R, Šlaus M, Novak M, Marjanović D. Y Chromosome Story-Ancient Genetic Data as a Supplementary Tool for the Analysis of Modern Croatian Genetic Pool. Genes (Basel). 2024 Jun 6;15(6):748. doi: 10.3390/genes15060748. PMID: 38927684; PMCID: PMC11202852 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC11202852/

Western hunter-gatherer, Wikipedia, This page was last edited on 18 August 2025, https://en.wikipedia.org/wiki/Western_hunter-gatherer

Posth, C., Yu, H., Ghalichi, A. et al. Palaeogenomics of Upper Palaeolithic to Neolithic European hunter-gatherers. Nature 615, 117–126 (2023). https://doi.org/10.1038/s41586-023-05726-0

[3] Primorac D, Šarac J, Havaš Auguštin D, Novokmet N, Bego T, Pinhasi R, Šlaus M, Novak M, Marjanović D. Y Chromosome Story-Ancient Genetic Data as a Supplementary Tool for the Analysis of Modern Croatian Genetic Pool. Genes (Basel). 2024 Jun 6;15(6):748. doi: 10.3390/genes15060748. PMID: 38927684; PMCID: PMC11202852. (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC11202852/

Szécsényi-Nagy Anna, et al. Tracing the genetic origin of Europe’s first farmers reveals insights into their social organization. Proc Biol Sci. 2015 Apr 22;282(1805):20150339. doi: 10.1098/rspb.2015.0339. PMID: 25808890; PMCID: PMC4389623 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC4389623/

Z. Hofmanová, S. Kreutzer, G. Hellenthal, C. Sell, Y. Diekmann, D. Díez-del-Molino, L. van Dorp, S. López, A. Kousathanas, V. Link, K. Kirsanow, L.M. Cassidy, R. Martiniano, M. Strobel, A. Scheu, K. Kotsakis, P. Halstead, S. Triantaphyllou, N. Kyparissi-Apostolika, […] & J. Burger, Early farmers from across Europe directly descended from Neolithic Aegeans, Proc. Natl. Acad. Sci. U.S.A.113 (25) 6886-6891, https://doi.org/10.1073/pnas.1523951113(2016).

[4] Hay, Maciamo, Haplogroups of Neolithic Europeans, Eupedia, https://www.eupedia.com/genetics/haplogroups_of_neolithic_farmers.shtml

Primorac D, Šarac J, Havaš Auguštin D, Novokmet N, Bego T, Pinhasi R, Šlaus M, Novak M, Marjanović D. Y Chromosome Story-Ancient Genetic Data as a Supplementary Tool for the Analysis of Modern Croatian Genetic Pool. Genes (Basel). 2024 Jun 6;15(6):748. doi: 10.3390/genes15060748. PMID: 38927684; PMCID: PMC11202852 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC11202852/

António Faustino Carvalho, Eva Fernández-Domínguez, Eduardo Arroyo-Pardo, Catherine Robinson, João Luís Cardoso, João Zilhão, Mário Varela Gomes, Hunter-gatherer genetic persistence at the onset of megalithism in western Iberia: New mitochondrial evidence from Mesolithic and Neolithic necropolises in central-southern Portugal, Quaternary International, Volumes 677–678, 2023, Pages 111-120, ISSN 1040-6182, https://doi.org/10.1016/j.quaint.2023.03.015 .
(https://www.sciencedirect.com/science/article/pii/S104061822300099X )

[5] The Chalcolithic period, or Copper Age, is the phase that followed the Neolithic (New Stone Age) and preceded the Bronze Age. It is characterized by the discovery and use of copper metallurgy, marking a transition from the exclusive use of stone tools to the use of early metal tools alongside existing stone technology.This innovation led to advancements in agriculture, crafts, trade, and societal complexity, setting the stage for the more advanced Bronze Age societies

Chalcolithic, Wikipedia, This page was last edited on 19 September 2025, https://en.wikipedia.org/wiki/Chalcolithic

References for the facts discussed in the paragraph:

Rivollat, M.; Jeong, C.; Schiffels, S.; Kücükkalıpcı, İ.; Pemonge, M.-H.; Rohrlach, A. B.; Alt, K. W.; Binder, D.; Friederich, S.; Ghesquière, E.; Gronenborn, D.; Laporte, L.; Lefranc, P.; Meller, H.; Réveillas, H.; Rosenstock, E.; Rottier, S.; Scarre, C.; Soler, L.; Wahl, J.; Krause, J.; Deguilloux, M.-F.; Haak, W.: Ancient genome-wide DNA from France highlights the complexity of interactions between Mesolithic hunter-gatherers and Neolithic farmers. Science Advances 6 (22), eaaz5344 , pp. 1 – 16 (2020), https://www.science.org/doi/10.1126/sciadv.aaz5344

Arzelier, Ana & Rivollat, Maïté & Harmony, De & Marie-Hélène, Pemonge & Binder, Didier & Convertini, Fabien & Henri, Duday & Gandelin, Muriel & Jean, Guilaine & Wolfgang, Haak & Deguilloux, Marie-France & Pruvost, Melanie. (2022). Neolithic genomic data from Southern France showcase intensified interactions with hunter-gatherer communities. iScience. 25. 105387. 10.1016/j.isci.2022.105387 , https://www.researchgate.net/publication/364419697_Neolithic_genomic_data_from_Southern_France_showcase_intensified_interactions_with_hunter-gatherer_communities

Andrew Zeilstra and Johanna Knop, Heightened Interaction Between Neolithic Migrants and Hunter-Gatherers in Western Europe, 29 May 2020 Press Release, Max Planck Institte of Geoanthropology, https://www.shh.mpg.de/1713184/haak-french-dna#_ftnref4

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 2025.03.24.644985; doi: https://doi.org/10.1101/2025.03.24.644985, https://www.biorxiv.org/content/10.1101/2025.03.24.644985v1.full; also found at: https://pubmed.ncbi.nlm.nih.gov/40196638

Tautalus, Geneticv Study: Long-term hunter-gatherer continuity in the Rhine-Meuse region was disrupted by local formation of expansive Bell Beaker groups, 25 Mar 2025, Forums > Population Genetics > Paleogenetics, Eupedia, https://www.eupedia.com/forum/threads/long-term-hunter-gatherer-continuity-in-the-rhine-meuse-region-was-disrupted-by-local-formation-of-expansive-bell-beaker-groups.45663/

Primorac D, Šarac J, Havaš Auguštin D, Novokmet N, Bego T, Pinhasi R, Šlaus M, Novak M, Marjanović D. Y Chromosome Story-Ancient Genetic Data as a Supplementary Tool for the Analysis of Modern Croatian Genetic Pool. Genes (Basel). 2024 Jun 6;15(6):748. doi: 10.3390/genes15060748. PMID: 38927684; PMCID: PMC11202852 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC11202852/

Szécsényi-Nagy Anna, et al. Tracing the genetic origin of Europe’s first farmers reveals insights into their social organization. Proc Biol Sci. 2015 Apr 22;282(1805):20150339. doi: 10.1098/rspb.2015.0339. PMID: 25808890; PMCID: PMC4389623 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC4389623/

Wolfgang Haak, Oleg Balanovsky, Juan J. Sanchez, Sergey Koshel, Valery Zaporozhchenko, Christina J. Adler, Clio S. I. Der Sarkissian, Guido Brandt, Carolin Schwarz, Nicole Nicklisch, Veit Dresely, Barbara Fritsch, Elena Balanovska, Richard Villems, Harald Meller, Kurt W. Alt, Alan Cooper, Ancient DNA from European Early Neolithic Farmers Reveals Their Near Eastern Affinities, PLOS Biology, 9 Nov 2010, https://doi.org/10.1371/journal.pbio.1000536 

[6] Source of quote:

Hay, Maciamo, The great pairings of Y-DNA haplogroups in prehistory, 19 Jul 2015, Forums > Population Genetics > Y-DNA Haplogroups, Eupedia, https://www.eupedia.com/forum/threads/the-great-pairings-of-y-dna-haplogroups-in-prehistory.31431/

See also the following for explaining the transition to farming between hunter gatherer and early farming groups:

Leendert P. Louwe Kooijmans,” Transition to Farming Along the Lower Rhine and Meuse .” Ancient Europe, 8000 B.C. to A.D. 1000: Encyclopedia of the Barbarian World. . Encyclopedia.com. (September 3, 2025). https://www.encyclopedia.com/humanities/encyclopedias-almanacs-transcripts-and-maps/transition-farming-along-lower-rhine-and-meuse

[7] Kamjan S, Gillis RE, Çakırlar C, Raemaekers DCM. Specialized cattle farming in the Neolithic Rhine-Meuse Delta: Results from zooarchaeological and stable isotope (δ18O, δ13C, δ15N) analyses. PLoS One. 2020 Oct 21;15(10):e0240464. doi: 10.1371/journal.pone.0240464. PMID: 33085689; PMCID: PMC7577484. (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC7577484/

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://pmc.ncbi.nlm.nih.gov/articles/PMC11974744/

“Transition to Farming Along the Lower Rhine and Meuse .” Ancient Europe, 8000 B.C. to A.D. 1000: Encyclopedia of the Barbarian World. . Retrieved September 03, 2025 from Encyclopedia.com: https://www.encyclopedia.com/humanities/encyclopedias-almanacs-transcripts-and-maps/transition-farming-along-lower-rhine-and-meuse

Kamjan S, Gillis RE, Çakırlar C, Raemaekers DCM (2020) Specialized cattle farming in the Neolithic Rhine-Meuse Delta: Results from zooarchaeological and stable isotope (δ¹⁸O, δ¹³C, δ¹⁵N) analyses. PLoS ONE 15(10): e0240464. https://doi.org/10.1371/journal.pone.0240464 

[8] 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 2025.03.24.644985; doi:  https://doi.org/10.1101/2025.03.24.644985, https://www.biorxiv.org/content/10.1101/2025.03.24.644985v1.full; also found at: https://pubmed.ncbi.nlm.nih.gov/40196638

Kivisild T. The study of human Y chromosome variation through ancient DNA. Hum Genet. 2017 May;136(5):529-546. doi: 10.1007/s00439-017-1773-z. Epub 2017 Mar 4. Erratum in: Hum Genet. 2018 Oct;137(10):863. doi: 10.1007/s00439-018-1937-5. PMID: 28260210; PMCID: PMC5418327 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC5418327/

Haak W, Lazaridis I, Patterson N, Rohland N, Mallick S, Llamas B, Brandt G, Nordenfelt S, Harney E, Stewardson K, Fu Q, Mittnik A, Bánffy E, Economou C, Francken M, Friederich S, Pena RG, Hallgren F, Khartanovich V, Khokhlov A, Kunst M, Kuznetsov P, Meller H, Mochalov O, Moiseyev V, Nicklisch N, Pichler SL, Risch R, Rojo Guerra MA, Roth C, Szécsényi-Nagy A, Wahl J, Meyer M, Krause J, Brown D, Anthony D, Cooper A, Alt KW, Reich D. Massive migration from the steppe was a source for Indo-European languages in Europe. Nature. 2015 Jun 11;522(7555):207-11. doi: 10.1038/nature14317. Epub 2015 Mar 2. PMID: 25731166; PMCID: PMC5048219 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC5048219/

[9] 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 2025.03.24.644985; doi: https://doi.org/10.1101/2025.03.24.644985, https://www.biorxiv.org/content/10.1101/2025.03.24.644985v1.full; also found at: https://pubmed.ncbi.nlm.nih.gov/40196638

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[13] da Silva, Marina Soares, Archaeogenetics of two subcontinents: the transition to Metal Ages in South Asia and Southwest Europe, PhD Thesis, Sep 2019, University of Huddlesfield, https://eprints.hud.ac.uk/id/eprint/35328/1/FINAL%20THESIS%20-%20Da%20Silva.pdf

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[14] Altena, Eveline, Smeding, Risha, van der Gaag, Kristaan J., de Leeuw, Rick,H., Vaske, Eileen, Reusink, Paul, Diekmann, Yoan, Thomas, Mark G., de Kniff, Peter, The dutch Y-chromosome from the early middle ages to present day. Archaeol Anthropol Sci 17, 116 (2025). https://doi.org/10.1007/s12520-025-02224-4

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[16] Altena, Eveline, et al., The dutch Y-chromosome from the early middle ages to present day. Archaeol Anthropol Sci 17, 116 (2025). https://doi.org/10.1007/s12520-025-02224-4

What Ancient DNA Reveals About the Medieval Population of the Low Countries, Mediievalist.net, https://www.medievalists.net/2025/05/ancient-dna-low-countries/

[17] 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. https://doi.org/10.1038/s41431-019-0496-0

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See also: 

Lucas G, Vésteinsson O. The Future of Periodization. Dissecting the Legacy of Culture History. Cambridge Archaeological Journal. 2024;34(4):637-652. doi: https://10.1017/S0959774324000015https://www.cambridge.org/core/journals/cambridge-archaeological-journal/article/abs/future-of-periodization-dissecting-the-legacy-of-culture-history/5CFAB2DAD8ACA218551D52B07F761049

Kotsonas, Antonis, Politics of Periodization and the Archaeology of Early Greece, Open Access on AJA Online, American Journal of Archaeology, Volume 120, Number 2, April 2016, Pages 239–70, DOI: 10.3764/aja.120.2.0239, https://www.journals.uchicago.edu/doi/pdf/10.3764/aja.120.2.0239

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Bevan A, Crema ER. 2021, Modifiable reporting unit problems and time
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Fiveable. “6.4 Limitations and Challenges in Archaeological Dating – Intro to Archaeology.” Fiveable, 2024. Accessed September 1, 2025. https://library.fiveable.me/introduction-archaeology/unit-6/limitations-challenges-archaeological-dating/study-guide/blc4ifVcgX2p9zp9 

Lucian George, Jade McGlyn, eds, Rethinking Period Boundaries: New Approaches to Continuity and Discontinuity in Modern European History and Culture, Oldenburg: De Gruyter, 2022

[23] Precise starting dates can fluctuate by a century or two, depending on the location within the watershed and archaeological definitions, as some areas may have experienced slightly later adoption of bronze technologies. However, the general consensus among recent regional studies places the Meuse-Rhine watershed Bronze Age onset in the early second millennium BCE.

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

Huth, Christoph, Water between two worlds – relections on the explanatory value of archaeological finds in a Bronze Age river landscape, in Ann Lehoërff and Marc Talon, eds, Movement Exchange and Identity in Europe in the Second and First Millennia BC, Oxford: Oxbrow Books, 2017 , 276-289, https://www.academia.edu/35831287/Water_between_two_worlds_reflections_on_the_explanatory_value_of_archaeological_finds_in_a_Bronze_Age_river_landscape

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, OliviaCheronet, Trudi Frost, Lora Iliev, Matthew Mah, Adam Micco, Jonas Oppenheimer, IrisPatterson, Lijun Qiu, Gregory Soos, J. Noah Workman, Ceiridwen J. Edwards, IosifLazaridis, Swapan Mallick, Nick Patterson, Nadin Rohland, Martin B. Richards, RonPinhasi, 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, https://www.biorxiv.org/content/10.1101/2025.03.24.644985v1.full

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W. K. van Zijverden, P.F.B. Jongste & F.S. Zuidhoff, Landscape and Occupation, Long Term Developments in the Dutch River Area During the Bronze Age, in De Dapper, Morgan, Frank Vermeulen, Sarah Deprez, and Devi Taelman, eds. 2009. “Ol’ Man River : Geo-Archaeological Aspects of Rivers and River Plains.” Ghent, Belgium: Academia Press. 618-627, https://www.academia.edu/1983135/LANDSCAPE_AND_OCCUPATION_LONG_TERM_DEVELOPMENTS_IN_THE_DUTCH_RIVER_AREA_DURING_THE_BRONZE_AGE

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Verhagen, Jan G.M., Sjoerd J. Kluiving, Emiel Anker, Liz van Leeuwen, Maarten A Prins, , Geoarchaeological prospection for Roman waterworks near the late Holocene Rhine-Waal delta bifurcation, the Netherlands, Catena, 2016, http://dx.doi.org/10.1016/j.catena.2016.03.027 , https://www.academia.edu/86389006/Geoarchaeological_prospection_for_Roman_waterworks_near_the_late_Holocene_Rhine_Waal_delta_bifurcation_the_Netherlands

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[29] Roymans, Nico and Stijn Heeren, Romano-Frankish interaction in the Lower Rhine frontier zone from the late 3rd to the 5th century – Some key archaeological trends explored, Germania, 99, 2021, pp. 133-156, https://doi.org/10.11588/ger.2021.92212

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See also:

Fokkens, Harry, and Anthony Harding, ‘Introduction: The Bronze Age of Europe’, in Harry Fokkens, and Anthony Harding (eds), The Oxford Handbook of the European Bronze Age (2013; online edn, Oxford Academic, 5 Sept. 2013), https://doi.org/10.1093/oxfordhb/9780199572861.013.0001, 

Stijn Arnoldussen and Harry Fokkens, Bronze Age Settlements in the Low Countries, 2008, Published by:  Oxbow Bookshttps://doi.org/10.2307/j.ctt1cfr8w0

[32] Fokkens, Harry, The Periodisation of the Dutch Bronze Age: a critical review, in Metz, W.H. Beek, B.L. van Steegstra, H (ed.), Essays presented to Jay Jordan Butler on the occasion of his 80th birthday (pp.241-262) Metz, Van Beek & Steegstra , 2001,  uploaded to ResearchGate by Harry Fokkens on 14 October 2015, https://www.researchgate.net/publication/28646850_The_periodisation_of_the_Dutch_Bronze_Age_a_critical_review

[33] Furholt, Martin. “Mobility and Social Change: Understanding the European Neolithic Period after the Archaeogenetic Revolution.” Journal of Archaeological Research 29, no. 4 (2021): 481–535, 637. https://www.jstor.org/stable/48771100 see also: https://www.semanticscholar.org/paper/Mobility-and-Social-Change:-Understanding-the-after-Furholt/a16b603d91b0533de47784d89e03693215589691

Furholt, Martin,  De-contaminating the aDNA–Archaeology Dialogue on Mobility and Migration Discussing the Culture-Historical Legacy, CURRENT SWEDISH ARCHAEOLOGY VOL. 27 2019 | https://doi.org/10.37718/CSA.2019.03

The Centre for Advanced Study, Dissecting the current debates on prehistoric migration, 7 Dec 2021, Press Release, partner.sciencenorway.no, https://partner.sciencenorway.no/archaeology-cas-centre-for-advanced-study-dna/dissecting-the-current-debates-on-prehistoric-migration/1944336

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Papac L, Ernée M, Dobeš M, Langová M, Rohrlach AB, Aron F, Neumann GU, Spyrou MA, Rohland N, Velemínský P, Kuna M, Brzobohatá H, Culleton B, Daněček D, Danielisová A, Dobisíková M, Hložek J, Kennett DJ, Klementová J, Kostka M, Krištuf P, Kuchařík M, Hlavová JK, Limburský P, Malyková D, Mattiello L, Pecinovská M, Petriščáková K, Průchová E, Stránská P, Smejtek L, Špaček J, Šumberová R, Švejcar O, Trefný M, Vávra M, Kolář J, Heyd V, Krause J, Pinhasi R, Reich D, Schiffels S, Haak W. Dynamic changes in genomic and social structures in third millennium BCE central Europe. Sci Adv. 2021 Aug 25;7(35):eabi6941. doi: 10.1126/sciadv.abi6941. PMID: 34433570; PMCID: PMC8386934. (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC8386934/

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[38] A “star-like phylogeny” is a genetic tree showing a central node from which many short branches radiate, indicating a recent and rapid demographic expansion from a common ancestral lineage. This pattern is a classic indicator of a founder effect, a form of genetic bottleneck. 

Ailsa Allaby and Michael Allby “star phylogeny .” A Dictionary of Earth Sciences. . Encyclopedia.com. 2 Sep. 2025 . https://www.encyclopedia.com/earth-and-environment/ecology-and-environmentalism/environmental-studies/star-phylogeny#:~:text=star%20phylogeny%20In%20a%20phylogenetic,Modern%20Language%20Association

Founder effect

The founder effect occurs when a new population is established by a very small number of individuals from a larger population. This results in a new population with: 

• Reduced genetic diversity: The founders carry only a fraction of the genetic variation present in the original population.
• Unique genetic composition: The new population’s gene pool is a non-random sample of the source population’s genes. Some rare alleles may be over-represented, while others may be lost entirely. 

Ramachandran, S. , Deshpande, O. , Roseman, C. C. , Rosenberg, N. A. , Feldman, M. W. , & Cavalli‐Sforza, L. L. (2005). Support from the relationship of genetic and geographic distance in human populations for a serial founder effect originating in Africa. Proceedings of the National Academy of Sciences, 102, 15942–15947. 10.1073/pnas.0507611102 (PubMed) https://pubmed.ncbi.nlm.nih.gov/16243969/

How a star-like phylogeny is formed

1. Founding event: A small group of individuals (the founders) with a limited gene pool moves to a new location.
2. Population expansion: The new population grows rapidly, possibly due to a lack of competitors or predators.
3. Low mutation accumulation: Since the population expanded very quickly, there has not been enough time for many new mutations to accumulate in the different lineages.
4. Short, radiating branches: The minimal genetic differences among individuals result in a phylogeny with many short branches emanating from a single, central common ancestor (the founder or founder lineage). 

The star-like pattern provides two key pieces of evidence for recent and rapid growth: 

• Short branches: The short length of the branches indicates that very few mutations occurred in each lineage since the founding event, suggesting that not much time has passed. 
• Common ancestor: All individuals can trace their ancestry back to a single, recent founder lineage, represented by the central node.