Looking at the Griff(is)(es)(ith) Y-DNA Phylogenetic Gap Associated with the Meuse and Rhine River Watershed – Part One

This story is the fourth part of a continuation of my focus on the G Haplogroup phylogenetic tree of the Griff(is)(es)(ith) patrilineal line of descent. The story discusses the migratory route of the Griffis family Y-DNA in the long term genealogical time layer.

This fourth part of the story focuses on examining possible macro social-cultural and enviromental influences that may explain the lack of identified subclades (ancestors) in the migratory path of the Griffis genetic paternal line. The period of time is roughly between 3000 BC and 650 CE.

The 2,850 year Gap between G-FGC7516 and G-Z6748: This phylogenetic gap was associated with haplogroup G-FGC477. 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.

Illustration one depicts the estimated migratory path of the Griff(is)(es)(ith) paternal genetic line. The first ‘phylogenetic gap’, depicted in the center of illustration one below, generally coincides with the migratory path of the Early European Farmers that followed the Dunube RIver watershed. Similar to the first phylogenetic gap that was discussed in a previous story, the second phylogenetic gap, depicted in the upper left hand portion of illustration one, follows another major European river watershed: the Meuse and Rhine River watersheds.

Illustration One: The Two Phylogenetic Gaps

Click for Larger View | Source: Migratory path of ancestors of G-Y132505, 10 Feb 2025, utilizing FamilyTreeDNA Globe Trekker

The Genetic Gap Between G-FGC7516 and G-Z6748

As reflected in table one below,, there are three most recent common genetic ancestors (MRCAs) associated with YDNA G haplogroups that are at the beginning of the second phylogenetic gap. They are estimated to have lived fairly close to each other in an area between the modern Gernman-Luxembourg border and the Mosel River (as depicted in illustration one above and five below). Their estimated dates of birth are between 2550 BCE and 2250 BCE.

Table One: G Haplogroups Associated with the Phylogenetic Gap

Click for Larger View | Source: Ancestral Path Chart for Haplogroup G-BY21678, FamilyTreeDNA, https://discover.familytreedna.com/y-dna/G-BY211678/path ; Y-DNA Haplotree – By SNP Variants for Confirmed Haplogroup is G-BY211678, FamilyTreeDNA, https://www.familytreedna.com/my/y-dna-haplotree

FamilyTreeDNA estimates indicate that the ancestor associated with the G-FGC7516 haplogroup was born around 2169 BCE. The level of certainty in determining the birth date is depicted in a probability plot in illustration two, which shows the most likely time when the common ancestor was born based on the statistical possibilities of various historical factors and data. It is ninety five percent certain that the ancestor associated with G-FGC7516 was born between 2892 BCE and 1551 BCE.

Illustration Two: Statistical Estimation Details on Haplogroup G-FGC7516

Click for Larger View | Source: FamilyTreeDNA,Scientific Details on Haplogroup G-FGC7516, https://discover.familytreedna.com/y-dna/G-FGC7516/scientific?section=tmrca

It has been estimated that the ancestor that marks the endpoint of this phylogenetic gap, who represents the G-Z6748 haplogroup, was born around 668 CE. There is a 95 percent chance of certainty that he was born between 380 – 908 CE (see illustration three).

Illustration Three: Statistical Estimation Details on Haplogroup G-G-Z6748

Click for Larger View | Source: FamilyTreeDNA,Scientific Details on Haplogroup G-Z6748, https://discover.familytreedna.com/y-dna/G-Z6748/scientific

As indicated above, there were two other haplogroups that were geographically and temporally close to G-FGC7516. The estimated ranges for the birthdates for the three genetic ancestors associated with these three haplogroups overlap.

Illustration Four: Overlapping Ranges of Estimated Birth Dates for G Haplogroups Associated with the Second Phylogenetic Gap

Click for Larger View

The estimated range, based on a 95 percent confidence interval, of when the ancestors associated with each of these haplogroups were born, is relatively large. As reflected in table two, given the emergence and shifting locations of various social groups and cultures during this period of time, these ancestors that were at the beginning of the gap could have been part of the Corded Ware or Bell Beaker cultures or possibly local bronze-Iron age tribes in the Meuse-Rhine watershed. The ancestor associated with the end of the gap could have been associated with Roman or post Roman transitions or German tribal societies. (see table two). The unknown ancestors in between these two time periods could have been part of various cultures in the Meuse-Rhine Watershed.

Table Two: Main Groups and Cultures Between 3000 BCE and 650 CE

PeriodMain Groups/CulturesKey Features
3000–2500 BCESwifterbant, Hazendonk, Vlaardingen, Seine-Oise-Marne (SOM) Groups, & Funnelbeaker Culture (TRB)Wetland hunter-gatherers, early farmers, high continuity
2500–1800 BCECorded Ware &
Bell Beaker Cultures
Steppe ancestry influx, major demographic/cultural shift
1800 BCE–First Century BCELocal Bronze/Iron Age tribes,
Urnfield culture (1300-750 BCE)
Hallstatt culture (800-450 BCE)
Persistence of distinct communities, gradual tribalization
First Century BCE – 650 CETexandri, Treveri, Triboci, Tubantes, Tungri, Tulingi, Germanic/Celtic tribes,
La Tène culture (450-50 BCE)
Tribal societies, Roman and post-Roman transitions, Frankish Expansion

If we consider the statistical variance in estimating the birth of the ancestors asociated with the endpoints of this phylogenetic gap, we are looking at a time span roughly between 2900 BCE and 908 CE. Prior to this phylogenetic time gap, it is possible that a Most Common Ancestor of the Griff(is)es)(ith) paternal line, associated with haplogroup G-Z1817, lived in the lower region of this watershed area between 3885 BCE and 2442 (see illustration five).

Illustration Five: Statistical Estimation Details on Haplogroup G-G-Z1817

Click for Larger View | Source: FamilyTreeDNA, Scientific Details of Haplogroup G-Z1817, https://discover.familytreedna.com/y-dna/G-Z1817/scientific

How the YDNA Haplgroup Map of the Meuse and Rhine Watershed Changed During This Time Frame

Between 2200 BCE and 650 CE, Northwestern Europe experienced dramatic shifts in Y-chromosome haplogroup distributions. These shifts were driven by migrations, cultural expansions, kinship practices, population replacements and environmental conditions (see table three).

At the beginning of this time frame, Early European Farmers (EEF), represented predominately by the YDNA G2a subclade interacted with the local Western Hunter-Gatherers (WHG), specifically YDNA subclades I2aR1b-V88, and C1a (typical of Mesolithic hunter-gatherers). Archeological remains of mitochondrial DNA showed predominant Neolithic farmer ancestry, indicating female mediated gene flow from farming communities. [1]

The common ancestors associated with the haplgroups G-Z1817, G-Z727, G-FGC477, and G-FGC7516 of the Griff(is)(es)(ith) line would have existed during this time frame.

The Bell Beaker culture (2500 – 1800 BCE) introduced the YDNA R1b-M269 and Rib-P312 subclades in Western Europe, eventually replacing about 90 percent of earlier Neolithic lineages. Between 2200 BCE and 650 CE, the G haplogroup lineages in the Rhine-Meuse watershed transitioned from Neolithic dominance to near-marginalization, making them demographic outliers compared to the influx of predominant steppe-derived lineages (R1b-M269 and R1a). Their near eradication reflects the region’s shift from Neolithic agrarian societies to Bronze Age pastoralist dominated populations. [2]

The Rhine-Meuse watershed served as a hub for the formation and expansion of Bell Beaker cultural groups, facilitating the spread of R1b subclades across Northwestern Europe. However, this region’s unique ecology allowed prolonged Western Hunter-Gatherers’ (WHG) genetic survival until Yamnaya pastoral Western Steppe Herder (WSH) male lineages achieved near-complete dominance by the Bronze Age.

Table Three: Key Genetic Shifts During the Phylogenetic Gap

PeriodDominant HaplogroupsCultural DriversGenetic Impact
Bronze Age
2200–800 BCE
G2a, I2a, R1b-V88 → R1b-U106/P312Bell Beaker,
Corded Ware
Admixture 40–90%
male replacement
Iron Age to Roman Period
800 – 400 CE
R1b subclades, I1Celtic / Germanic regionalizationLocal diversification
Germanic Expansions
400–650 CE
R1b-U106, R1b-P312, R1a-M420, I1Anglo-Saxon
migrations
73–86% turnover
in Britain

These shifts reflect Northwestern Europe’s complex interplay of steppe ancestry expansions, later Germanic tribal migrations, and cultural hybridization over three millennia.

Limited adoption of Corded Ware pottery culture (around 2500 BCE) was introduced through haplogroup R1b-U106 (a steppe-associated lineage) to the region. However, autosomal steppe ancestry remained low (around 11 percent), suggesting cultural adoption without large-scale population replacement. [3]

Between 2500 BCE and 1800 BCE, the Bell Beaker culture (2500–1800 BCE) introduced the R1b-M269 subclades (particularly R1b-P312) to Western Europe, replacing about ninety percent of earlier Neolithic Y-lineages[4] This lineage became dominant in Britain, France, and Iberia, correlating with steppe ancestry. [5] The Bell Beaker complex marked a 83 to 91 percent genetic turnover, blending R1b-U106 and R1b-P312 lineages (from Corded Ware migrants) with 9 to 17 percent local Rhine-Meuse ancestry (high in hunter-gatherer ancestry).

In Central Europe, Corded Ware groups spread R1a-M417 (ancestral to R1a-Z645) from the Pontic-Caspian steppe, dominating regions like Bohemia and Scandinavia. [6] A steppe ancestry surge, reflecting a forty to fifty percent genetic replacement, occurred in Britain and Iberia, with R1b-P312 reaching 70–90% frequency among males. [7] While some earlier G lineages declined, certain subclades of G, such as G-L13, may have arrived with the steppe pastoralists, and their descendants. This highlights the complex nature of human migration and the subsequent mixing of populations and genetic lineages throughout European history. [8]

The ancestors of the Griff(is)(es)(ith) paternal genetic lineage that migrated through this area during this time period became genetic outliers among other predominant genetic lineages in this geographical area (see table four).

Table Four: G haplogroup’s Genetic Legacy

Time Period
within
Phylogenetic Gap
G Haplogroup Status
within Region
Key Drivers
Neolithic
(G-Z1817, G-Z727,
G-FGC477, G-FGC7516)
Dominant (~60% of males)Agricultural expansion
Bronze/ Iron Age & Roman Era
(Absence of Phylogenetic Subclades)
Near eradication (<5%)Steppe migrations
Germanic Expansions
(Absence of Subclades &
Endpoint of G-Z6748)
Marginal (1–2% in NW Europe)Continued admixture

By 650 CE, the G haplogroup represented less that two percent of YDNA lineages in Northwestern Europe, its decline emblematic of the region’s shift from Neolithic agrarian societies to Bronze/Iron Age pastoralist-dominated populations. [9]

The Uniqueness of the Meuse and Rhine River Watershed

Illustration five depicts the estimated migratory path between haplogroup G-FGC716 and haplogroup G-Z6748, the two endpoints associated with this second phylogenetic gap. The map is part of a screen shot utilizing FamilyTreeDNA Globe Trekker program that maps out the YDNA migratory parth for the Griff(is)(es)(ith) genetic paternal line. The ‘zone of confidence’ reflected in the map depicts the possible variation of the migratiory path.

What is noteworthy when looking at illustration five is the relatively short distance between the estimated locations of the two ancestors that represent the endpoints of the genetic gap. The approximate locations of the two endpoints are Aachen, Germany and Amsterdam, Netherlands. Based on contemporary measures of distance, the length of this migratory gap is about 230 kilometers or 143 miles. Despite a relatively short distance between these two ancient ancestors, there are roughly 2,850 years between them.

Illustration Five: The Phylogenetic Gap between G-FGC7516 and G-Z6748

Click for Larger View | Source: Migratory path of ancestors of G-Y132505, 10 Feb 2025, utilizing FamilyTreeDNA Globe Trekker

The migratory path of the Griff(is)(es)(ith) paternal line follows the contours between the Rhine River and the Meuse River watershed (see illustration six). The Meuse and Rhine watershed played a critical role in shaping migration, cultural dynamics and the Y-DNA genetic mix in northwestern Europe. It is within this geographical context that we find the second major phylogenetic gap in the Griff(is)(es)(ith) migratory path. I have added circled areas in illustration six to depict the general, estimated locations of the endpoints of the phylogenetic gap.

Illustration Six: The Water Basins of the Rhine River and Meuse River and the Two Most Common Recent Ancestor Enpoints for the Phylogenetic Gap in the Context of Modern Day Poltical Boundaries

Click for Larger View | Source: Pilarczyk, Krystian. (2007). NATO Science Series. 10.1007/978-1-4020-5741-0_26. ,Fig. 1 The Rhine and Meuse basin Page 26 ,https://www.researchgate.net/figure/The-Rhine-and-Meuse-basin_fig11_226598873

The Rhine River and the Meuse River are part of one of northwest Europe’s significant watersheds. The rivers and related tribularies have served as a frontier, a cultural crossroad, a barrier between groups and a boundary between societies throughout human history. Archaeological evidence and historical records reveal a rich tapestry of cultures that developed along its banks over nearly three millennia. The Rhine and Meuse River regions between 2200 BCE and 650 CE hosted diverse cultures and societies shaped by Neolithic, Celtic, Germanic, and Roman influences, evolving from tribal networks to fortified Roman frontiers and later Frankish kingdoms.

The Rhine-Meuse delta’s dynamic wetland and riverine environments profoundly shaped the interaction of social groups and cultures from 3000 BCE to 800 CE, fostering adaptive subsistence strategies, trade networks, and the interaction of various cultures while also necessitating technological innovation to manage environmental challenges of flooding and changing contours of the land. The shifts in land types are reflected in the following video through time (Click on image to start the video. Video will play directly within the webpage on mobile browsers, instead of opening in a fullscreen player.)

Video: Historical Changes to the Landscape: 5500 BCE, 3850 BCE, 2750 BCE, 500 BCE and 50 BCE

Source: Historical Changes to Netherlands Lanscape 550 BCE to 50 BCE, Prehistory of the Netherlands, Updated, 15 May 2025, Wikipedia, https://en.wikipedia.org/wiki/Prehistory_of_the_Netherlands

The Rhine-Meuse delta’s dynamic ecology played a dual role in shaping Y haplogroup expansions, both hindering certain genetic lineages and promoting others through environmental strictures and localized adaptations.

The delta’s ecology primarily hindered the expansion of outside incoming YDNA haplogroups (e.g., steppe R1a, Neolithic G2a) by creating barriers to mass male migrations. Simultaneously, it promoted the persistence and of indigenous lineages through isolation, localized resource strategies, and unique admixture events. This dual dynamic produced a YDNA landscape distinct from neighboring European regions. [10]

The delta acted as both a conservator of ancient YDNA (until ~2500 BCE) and a launchpad for R1b lineages that reshaped European paternal ancestry. These patterns underscore how mobility strategies—pioneer groups, kinship networks, and riverine trade—could disproportionately propagate specific male lineages despite limited autosomal impact.

The Meuse and Rhine River watershed may have had a major influence on the Griff(is)(es)(ith) YDNA phylogenetic tree through dynamic interactions between ecological barriers, R haplogroup male-biased migrations, the locus of geopolitical conflicts and changing political boundaries.

The ecology of the Rhine-Meuse delta was characterized by dynamic wetlands, shifting river channels, and fragmented landscapes. This environment had a significant impact on the spread and unique evolution of YDNA haplogroups over time. [11]

YDNA Preservation Challenges in the Watershed Area

Optimal Y-DNA preservation occurred in cool, alkaline, waterlogged environments with rapid sedimentation, while warmer climates, acidic soils, and hydrological disturbances degraded genetic material. Human activities both aided (burial under sediment) and hindered (soil disturbance) the preservation of YDNA remains. [12]

Table Five: Ecological Watershed Conditions that Impact YDNA Retrieval

FactorsResultant Conditions
Hydrological & Sendimentary
Dynamics
Increased sedimentation from Bronze Age agriculture (~3000 BP) accelerated delta growth, burying and potentially preserving organic materials in anaerobic floodplain deposits [13]
Flooding phases (e.g., Roman Period, 12 BCE–250 CE) redistributed sediments, alternately exposing or sealing archaeological contexts [14]
Avulsion processes (channel shifts) created localized burial environments, with some areas like abandoned channels acting as natural preservation traps [15]
Climate & Microenvironmental
Factors
Cooler, drier periods (e.g., Subboreal, ~2800 BCE) slowed DNA degradation by reducing hydrolytic damage [16]
Alkaline environments (e.g., salt-rich deposits) inhibited DNA fragmentation by limiting water activity and enzymatic decay [17]
Anoxic conditions in waterlogged soils (e.g., deltaic peat) reduced microbial activity, preserving skeletal remains [18]
Human ImpactDeforestation and land clearance increased erosion, rapidly burying remains under sediment layers [19]
Roman-era mining and medieval embankments disturbed archaeological layers but also created sealed contexts (e.g., ditches) conducive to preservation [20]
Settlement patterns: Upland sites on sandy soils experienced higher DNA degradation due to acidic conditions, while delta wetlands offered better preservation [21]
Climate & Soil
Conditions
Warmer phases (e.g., Roman Warm Period) accelerated DNA fragmentation via increased microbial and enzymatic activity [22]
Cooler intervals (e.g., Late Antiquity cooling) enhanced preservation, particularly in deep sediment [23]
Soil chemistry: Bones in alkaline soils (pH >7) retained longer DNA fragments than those in acidic environments [24]

Ecological Barriers and Isolation

The delta’s constantly changing wetlands, river courses, and peat bogs created numerous ecological “islands.” These natural barriers limited large-scale migration and promoted the isolation of local populations. As a result, indigenous YDNA haplogroups—especially those associated with Mesolithic hunter-gatherers (e.g., Haplogroups I2a, R1b-V88, and C1a) persisted in the region much longer than in more accessible parts of Europe. This isolation fostered genetic drift within small, localized communities, leading to the retention and sometimes unique evolution of specific YDNA lineages. [25]

Selective YDNA Gene Flow and Sex-Based Admixture

The challenging terrain and waterlogged soils made the delta less attractive or accessible to large-scale male-dominated migrations, such as those associated with the G haplgroup based Neolithic farmers or the R haplgrop based later steppe populations. However, there was significant female-mediated gene flow, as evidenced by the predominance of Neolithic farmer mitochondrial (mtDNA) lineages alongside persistent local YDNA haplogroups.

This resulted in a unique genetic structure in the neolithic era: local men maintained hunter-gatherer YDNA, while women from incoming farming populations contributed new mtDNA lineages. This pattern is directly linked to the delta’s ecology, which favored gradual, piecemeal integration over wholesale population replacement. [26]

Buffering and Filtering of Major YDNA Migrations

The spread of Steppe ancestry (e.g., Corded Ware culture-associated R1a haplogroups) and later Bell Beaker migrations) was muted or filtered in the delta. While some Steppe and Bell Beaker YDNA lineages did appear, the dominant pattern was one of fusion with the local population rather than replacement. The delta’s fragmented, water-rich environment acted as a semi-permeable barrier, allowing for some genetic and cultural exchange but preserving a strong local genetic signature. [27]

Small Community Size and Kinship Structures

Genetic analyses show evidence of close kinship ties within local communities, consistent with small, relatively isolated populations. This would have reinforced the persistence of local YDNA haplogroups and limited the genetic impact of newcomers. [28]

Legacy of Ecological and Territorial Dynamics

These ecological effects created a legacy of territorial dynamics, as depicted in table four, that had subsequent effects through this phylogeneic gap period. The Rhine-Meuse delta’s ecological and associated cultural transformations between 2500 BCE and 600 CE significantly shaped the structure of individual YDNA phylogenetic trees through this time period..

Table Six: Genetic Legacy of Meuse-Rhine Ecological and Territorial Dynamics

Ecological
& Territorial Dynamics
in the
Phylogenetic
Gap
Y-Haplogroup ImpactExample Regions and Description
Neolithic and Iron Age Riverine Barriers [29] Preserved I2a and C1a in wetlands; restricted G2a and  R1b to elevated terraces Delta’s fragmented landscapes limited mass migrations, favoring small, kin-based groups. This preserved indigenous YDNA (e.g., I2a) until Bell Beaker groups leveraged river networks for rapid dispersal
Bronze Age Pastoralist Expansion [30]The Bronze Age brought a sex-biased migration pattern, with male Steppe lineages largely replacing previous Y-DNA profiles while incorporating some local female ancestry. This contrasts with earlier Neolithic transitions that showed more gradual admixture.Western/central Netherlands: Mixed subsistence strategy of hunting, gathering, and farming persisted until the 3rd millennium BCE, when a more intense farming-based economy emerged in association with the Late Vlaardingen complex and the introduction of the ard plough around 3000 BCE
Bronze Age Deforestation ~ 1500 BCE [31] Predominant Y-DNA haplogroups were R1b-L151-P312 subclades associated with Bell Beaker and Corded Ware cultures, while secondary lineages included remnants of earlier Neolithic farmer ancestry (G2a2 and I2a)Deforestation upstream increased sediment flow, reshaping the delta’s river network. This allowed new branches to form, facilitating trade and migration movement but also requiring flood management.
Roman Frontier Militarization [32]North of the Rhine : R1b-U106 enriched at Roman military camps or stations along the Rhine River); 

South/west of the Rhine: Higher frequencies of R1b-P312 (Celtic-associated) and residual G2a (Neolithic farmer) in civilian Roman controlled hinterlands

Celtic La Tène culture influenced southern delta groups, while Germanic tribes (e.g., Frisii, Batavi) occupied the north. The Rhine served as a cultural and political boundary

Rhine Limes as a Roman frontier region, facilitated significant migration, mixing, and demographic changes, potentially impacting the distribution and growth of various haplogroups during Roman times

Forts and towns were built on undercut river meanders with stabilized banks (e.g., Packwerk structures at Xanten) to ensure year-round harborage.

The Limes (Roman frontier) transitioned from a defensive line to a trade corridor by the 3rd century CE, guarded via mobile patrols as river activity increased
Post-Roman Shifts and Migrations [33] R1b-S116 replaced R1b-P312 in Frankish-controlled Meuse Valley; reinforced existing R1b-U106 dominance

Late Roman collapse: Post-3rd century CE Germanic incursions (e.g., Franks, Visigoths) introduced I1a and R1b-S116 subclades, replacing Roman-era lineages in contested zones like the Lower Rhine
Maastricht-Aachen corridor [21];
Renewed river dynamism after 300 CE isolated communities but reinforced existing R1b-U106 dominance, as these lineages were already adapted to floodplain management
Early Medieval Period (500–900 CE): Frankish Expansion and Male-Lineage Dominance [34] The Frankish expansion and territorial control from the 5th century CE amplified R1b-U106’s prevalence, as this group established political dominance in the region.

Rhineland refugia: Earlier Bell Beaker migrations (2500–1800 BCE) had established R1b-L151 lineages (including P312 and U106), which remained foundational; and retained older R1b-L51 and J2a haplogroups, linked to Roman-era military settlements

G2a subclades, minor Neolithic remnant lineages found along the Rhine-Meuse-Thames corridor, representing ~5% of male lineages


Meuse Valley: Environmental factors like the delta’s navigable waterways facilitated both trade networks and military campaigns, enabling sustained genetic inflow from Central Europe

Isolated communities in the Ardennes and Eifel regions [35]

Each of these ecological and territorial influences may have played a part in the lack of discovered subclades in the phylogenetic gap over an estimated 95 generations.

The Continuation – Part Two

Part two of this story focuses on the emergence and shifting locations of various social groups and cultures in the Muese Rhine watershed during this period of time. The ancestors that are documented as well as the undocumented generations may have been part of the cultures that lived in the Meuse-Rhine watershed.

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 GlobetrekkerTM 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] A significant cultural and demographic shift that occurred in some regions during the transition from the Neolithic to the Bronze Age. This shift involved a decline of sedentary farming communities and a rise of pastoralist groups, characterized by mobile livestock herding. 

The Neolithic period saw the development of agriculture and the establishment of permanent settlements. These societies were based on cultivating crops and raising domesticated animals, allowing for larger and more stable populations than hunter-gatherer societies.

The Bronze Age saw further technological advancements, including the development of bronze tools and weapons, as well as innovations like the wheel and horse domestication. These innovations facilitated the rise of pastoralist cultures, especially in the Eurasian steppes. These groups were highly mobile, relying on large herds of livestock, particularly horses, for sustenance and transport.

The shift to a pastoralist-dominated society might have led to the “near-eradication” of some Neolithic agrarian populations in certain regions. This could have occurred through various mechanisms, including:

  • Migration and Displacement: Mobile pastoralist groups could have migrated into areas occupied by farmers, displacing them and taking over their lands.
  • Conflict and Conquest: The increased mobility and military capabilities of pastoralists, enhanced by horse domestication, may have allowed them to conquer settled agrarian communities.
  • Disease: The movement of people and animals could have introduced new diseases to which settled populations had little immunity.
  • Cultural Assimilation: Over time, some agrarian populations may have adopted the pastoralist lifestyle and integrated into their societies. 

Archaeological and genetic studies support the idea of large-scale population shifts during this period, particularly in Europe. For example, the expansion of the Yamnaya culture, a Bronze Age pastoralist society from the Pontic-Caspian Steppe, into Northern Europe shows evidence of significant population replacement and admixture with existing hunter-gatherer and early farmer populations. 

Taylor, W.T.T., Clark, J., Bayarsaikhan, J. et al. Early Pastoral Economies and Herding Transitions in Eastern Eurasia. Sci Rep 10, 1001 (2020). https://doi.org/10.1038/s41598-020-57735-y

Neolithic Europe, Wikipedia, This page was last edited on 25 May 2025, https://en.wikipedia.org/wiki/Neolithic_Europe

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

Lasse Sørensen, Sabine Karg, The expansion of agrarian societies towards the north – new evidence for agriculture during the Mesolithic/Neolithic transition in Southern Scandinavia,
Journal of Archaeological Science, Volume 51, 2014, Pages 98-114,
https://doi.org/10.1016/j.jas.2012.08.042 .
(https://www.sciencedirect.com/science/article/pii/S0305440312003962 )

Ghalichi, A., Reinhold, S., Rohrlach, A.B. et al. The rise and transformation of Bronze Age pastoralists in the Caucasus. Nature 635, 917–925 (2024). https://doi.org/10.1038/s41586-024-08113-5

C. Jeong, et al, Bronze Age population dynamics and the rise of dairy pastoralism on the eastern Eurasian steppe, Proc. Natl. Acad. Sci. U.S.A. 115 (48) E11248-E11255 ,https://doi.org/10.1073/pnas.1813608115 (2018).

Robert Spengler , Michael Frachetti , Paula Doumani , Lynne Rouse , Barbara Cerasetti , Elissa Bullion and Alexei Mar’yashev, Early Agriculture and crop transmission among Bronze Age Mobile pastoralists of central Eurasia, 22 May 2014, Proceedings of the Royal society B, vol 281, Issue 1783  https://doi.org/10.1098/rspb.2013.3382

[2] 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, 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, Altena E, Bourgeois Q, Fokkens H, Amkreutz L, Deguilloux MF, Fichera A, Flas D, Gandini F, Kegler JF, Kootker LM, Leijnse K, Kooijmans LL, Lauwerier R, Miller R, Molthof H, Noiret P, Raemaekers DCM, Rivollat M, Smits L, Stewart JR, Anscher TT, Toussaint M, Callan K, Cheronet O, Frost T, Iliev L, Mah M, Micco A, Oppenheimer J, Patterson I, Qiu L, Soos G, Workman JN, Edwards CJ, Lazaridis I, Mallick S, Patterson N, Rohland N, Richards MB, Pinhasi R, Haak W, Pala M, Reich D. 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

Hay, Maciamo, Genetic History of the Benelux & France, Aug 2021, Eupedia, https://www.eupedia.com/europe/benelux_france_dna_project.shtml

Haplogroup G-M201, This page was last edited on 25 May 2025, Wikipedia, https://en.wikipedia.org/wiki/Haplogroup_G-M201

[3] 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/

[4] García-Fernández, C., Lizano, E., Telford, M. et al. Y-chromosome target enrichment reveals rapid expansion of haplogroup R1b-DF27 in Iberia during the Bronze Age transition.Sci Rep 12, 20708 (2022). https://doi.org/10.1038/s41598-022-25200-7

[5] Gretzinger J, Sayer D, Justeau P, Altena E, Pala M, Dulias K, Edwards CJ, Jodoin S, Lacher L, Sabin S, Vågene ÅJ, Haak W, Ebenesersdóttir SS, Moore KHS, Radzeviciute R, Schmidt K, Brace S, Bager MA, Patterson N, Papac L, Broomandkhoshbacht N, Callan K, Harney É, Iliev L, Lawson AM, Michel M, Stewardson K, Zalzala F, Rohland N, Kappelhoff-Beckmann S, Both F, Winger D, Neumann D, Saalow L, Krabath S, Beckett S, Van Twest M, Faulkner N, Read C, Barton T, Caruth J, Hines J, Krause-Kyora B, Warnke U, Schuenemann VJ, Barnes I, Dahlström H, Clausen JJ, Richardson A, Popescu E, Dodwell N, Ladd S, Phillips T, Mortimer R, Sayer F, Swales D, Stewart A, Powlesland D, Kenyon R, Ladle L, Peek C, Grefen-Peters S, Ponce P, Daniels R, Spall C, Woolcock J, Jones AM, Roberts AV, Symmons R, Rawden AC, Cooper A, Bos KI, Booth T, Schroeder H, Thomas MG, Helgason A, Richards MB, Reich D, Krause J, Schiffels S. The Anglo-Saxon migration and the formation of the early English gene pool. Nature. 2022 Oct;610(7930):112-119. doi: 10.1038/s41586-022-05247-2. Epub 2022 Sep 21. Erratum in: Nature. 2022 Nov;611(7934):E3. doi: 10.1038/s41586-022-05429-y. PMID: 36131019; PMCID: PMC9534755,( PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC9534755/

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/

Hay, Maciamo, Genetic History of he Benelux & France, Aug 2021, Eupedia, https://www.eupedia.com/europe/benelux_france_dna_project.shtml

Haplogroup G-M201, This page was last edited on 25 May 2025, Wikipedia, https://en.wikipedia.org/wiki/Haplogroup_G-M201

[6] Oğuzhan Parasayan et al., Late Neolithic collective burial reveals admixture dynamics during the third millennium BCE and the shaping of the European genome.Sci. Adv.10,eadl2468(2024).DOI:10.1126/sciadv.adl2468

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/

[7] Gretzinger J, Sayer D, Justeau P, Altena E, Pala M, Dulias K, Edwards CJ, Jodoin S, Lacher L, Sabin S, Vågene ÅJ, Haak W, Ebenesersdóttir SS, Moore KHS, Radzeviciute R, Schmidt K, Brace S, Bager MA, Patterson N, Papac L, Broomandkhoshbacht N, Callan K, Harney É, Iliev L, Lawson AM, Michel M, Stewardson K, Zalzala F, Rohland N, Kappelhoff-Beckmann S, Both F, Winger D, Neumann D, Saalow L, Krabath S, Beckett S, Van Twest M, Faulkner N, Read C, Barton T, Caruth J, Hines J, Krause-Kyora B, Warnke U, Schuenemann VJ, Barnes I, Dahlström H, Clausen JJ, Richardson A, Popescu E, Dodwell N, Ladd S, Phillips T, Mortimer R, Sayer F, Swales D, Stewart A, Powlesland D, Kenyon R, Ladle L, Peek C, Grefen-Peters S, Ponce P, Daniels R, Spall C, Woolcock J, Jones AM, Roberts AV, Symmons R, Rawden AC, Cooper A, Bos KI, Booth T, Schroeder H, Thomas MG, Helgason A, Richards MB, Reich D, Krause J, Schiffels S. The Anglo-Saxon migration and the formation of the early English gene pool. Nature. 2022 Oct;610(7930):112-119. doi: 10.1038/s41586-022-05247-2. Epub 2022 Sep 21. Erratum in: Nature. 2022 Nov;611(7934):E3. doi: 10.1038/s41586-022-05429-y. PMID: 36131019; PMCID: PMC9534755 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC9534755/

García-Fernández, C., Lizano, E., Telford, M. et al. Y-chromosome target enrichment reveals rapid expansion of haplogroup R1b-DF27 in Iberia during the Bronze Age transition.Sci Rep 12, 20708 (2022). https://doi.org/10.1038/s41598-022-25200-7

[8] Eugène C.W.L. (Boed) Marres, G-M201, Marres, https://www.marres.nl/EN/G-M201.htm

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

Haplogroup G-M201, Wikipedia, This page was last edited on 25 May 2025, https://en.wikipedia.org/wiki/Haplogroup_G-M201

[10]  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 Iñigo, 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/

[11] See for example:

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

Pierrik, H.J., E. Stouthamer, K.M. Cohen, Natural levee evolution in the Rhine-Meuse delta, theNetherlands, during the first millennium CE, Geomorphology, 295 (2017) 215-234, https://dspace.library.uu.nl/bitstream/handle/1874/352044/Natural.pdf?sequence=1

Arnoldussen, Stijn, A Living Landscape: Bronze Age Settlement Sites in the Dutch River area (c.2000-80 BC), Sidestone Press, 2008 https://www.sidestone.com/openaccess/9789088900105.pdf

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

Kooijmans, L.P. Louwe, The Rhine/Meuse Delta: Four Studies on its Prehistoric Occupation and Holecene Geology, Lieden University Press, 1974

Parasayan O, Laurelut C, Bôle C, Bonnabel L, Corona A, Domenech-Jaulneau C, Paresys C, Richard I, Grange T, Geigl EM. Late Neolithic collective burial reveals admixture dynamics during the third millennium BCE and the shaping of the European genome. Sci Adv. 2024 Jun 21;10(25):eadl2468. doi: 10.1126/sciadv.adl2468. Epub 2024 Jun 19. PMID: 38896620; PMCID: PMC1118650 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC11186501/

van Dinter, M., The Roman Limes in the Netherlands: how a delta landscape determined the location of the military structures, Netherlands Journal of Geosciences, 92-1, 11-13 2013, 11-32, https://www.researchgate.net/publication/287527529_The_Roman_Limes_in_the_Netherlands_How_a_delta_landscape_determined_the_location_of_the_military_structures

[12] Olson, K. , Krug, E. and Chernyanskii, S. (2025) Natural and Anthropic Environmental Risks to the Rhine River and Delta. Open Journal of Soil Science15, 235-267. doi: 10.4236/ojss.2025.154012.

Kravanja P, Golob A, Concato M, Leskovar T, Zupanič Pajnič I. Effects of different environmental factors on preservation of DNA in petrous bones: A comparative study of two Slovenian archaeological sites. Forensic Sci Int. 2025 Jun;371:112495. doi: 10.1016/j.forsciint.2025.112495. Epub 2025 May 7. PMID: 40349398 (PubMed) https://pubmed.ncbi.nlm.nih.gov/40349398/

Poetsch M, Markwerth P, Konrad H, Bajanowski T, Helmus J. About the influence of environmental factors on the persistence of DNA – a long-term study. Int J Legal Med. 2022 May;136(3):687-693. doi: 10.1007/s00414-022-02800-6. Epub 2022 Feb 23. PMID: 35195781; PMCID: PMC9005405 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC9005405/

Conor Rossi , Gabriela Ruß-Popa , Valeria Mattiangeli , Fionnuala McDaid , Andrew J. Hare , Hossein Davoudi , Haeedeh Laleh , Zahra Lorzadeh , Roya Khazaeli , Homa Fathi , Matthew D. Teasdale , Abolfazl A’ali , Thomas Stöllner, et al , Exceptional ancient DNA preservation and fibre remains of a Sasanian saltmine sheep mummy in Chehrābād, Iran, Biology Letter, The Royal Society, Jul 2021, 17, 7, https://royalsocietypublishing.org/doi/10.1098/rsbl.2021.0222

[13] Olson, K. , Krug, E. and Chernyanskii, S. (2025) Natural and Anthropic Environmental Risks to the Rhine River and Delta. Open Journal of Soil Science15, 235-267. doi: 10.4236/ojss.2025.154012.

[14] Peng, F., Prins, M. A., Kasse, C., Cohen, K. M., Van der Putten, N., van der Lubbe, J., Toonen, W. H. J., & van Balen, R. T. (2019). An improved method for paleoflood reconstruction and flooding phase identification, applied to the Meuse River in the Netherlands. Global and Planetary Change177, 213-224. https://doi.org/10.1016/j.gloplacha.2019.04.006

[15] Peng, F., Prins, M. A., Kasse, C., Cohen, K. M., Van der Putten, N., van der Lubbe, J., Toonen, W. H. J., & van Balen, R. T. (2019). An improved method for paleoflood reconstruction and flooding phase identification, applied to the Meuse River in the Netherlands. Global and Planetary Change177, 213-224. https://doi.org/10.1016/j.gloplacha.2019.04.006

Olson, K. , Krug, E. and Chernyanskii, S. (2025) Natural and Anthropic Environmental Risks to the Rhine River and Delta. Open Journal of Soil Science15, 235-267. doi: 10.4236/ojss.2025.154012.

[16] Logan Kistler, Roselyn Ware, Oliver Smith, Matthew Collins, Robin G. Allaby, A new model for ancient DNA decay based on paleogenomic meta-analysis, Nucleic Acids Research, Volume 45, Issue 11, 20 June 2017, Pages 6310–6320, https://doi.org/10.1093/nar/gkx361

Peng, F., Prins, M. A., Kasse, C., Cohen, K. M., Van der Putten, N., van der Lubbe, J., Toonen, W. H. J., & van Balen, R. T. (2019). An improved method for paleoflood reconstruction and flooding phase identification, applied to the Meuse River in the Netherlands. Global and Planetary Change177, 213-224. https://doi.org/10.1016/j.gloplacha.2019.04.006

Conor Rossi , Gabriela Ruß-Popa , Valeria Mattiangeli , Fionnuala McDaid , Andrew J. Hare , Hossein Davoudi , Haeedeh Laleh , Zahra Lorzadeh , Roya Khazaeli , Homa Fathi , Matthew D. Teasdale , Abolfazl A’ali , Thomas Stöllner, et al , Exceptional ancient DNA preservation and fibre remains of a Sasanian saltmine sheep mummy in Chehrābād, Iran, Biology Letter, The Royal Society, Jul 2021, 17, 7, https://royalsocietypublishing.org/doi/10.1098/rsbl.2021.0222

[17] Conor Rossi , Gabriela Ruß-Popa , Valeria Mattiangeli , Fionnuala McDaid , Andrew J. Hare , Hossein Davoudi , Haeedeh Laleh , Zahra Lorzadeh , Roya Khazaeli , Homa Fathi , Matthew D. Teasdale , Abolfazl A’ali , Thomas Stöllner, et al , Exceptional ancient DNA preservation and fibre remains of a Sasanian saltmine sheep mummy in Chehrābād, Iran, Biology Letter, The Royal Society, Jul 2021, 17, 7, https://royalsocietypublishing.org/doi/10.1098/rsbl.2021.0222

[18] Conor Rossi , Gabriela Ruß-Popa , Valeria Mattiangeli , Fionnuala McDaid , Andrew J. Hare , Hossein Davoudi , Haeedeh Laleh , Zahra Lorzadeh , Roya Khazaeli , Homa Fathi , Matthew D. Teasdale , Abolfazl A’ali , Thomas Stöllner, et al , Exceptional ancient DNA preservation and fibre remains of a Sasanian saltmine sheep mummy in Chehrābād, Iran, Biology Letter, The Royal Society, Jul 2021, 17, 7, https://royalsocietypublishing.org/doi/10.1098/rsbl.2021.0222

Kravanja P, Golob A, Concato M, Leskovar T, Zupanič Pajnič I. Effects of different environmental factors on preservation of DNA in petrous bones: A comparative study of two Slovenian archaeological sites. Forensic Sci Int. 2025 Jun;371:112495. doi: 10.1016/j.forsciint.2025.112495. Epub 2025 May 7. PMID: 40349398. (PubMed) https://pubmed.ncbi.nlm.nih.gov/40349398/

[19] “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. (June 17, 2025). https://www.encyclopedia.com/humanities/encyclopedias-almanacs-transcripts-and-maps/transition-farming-along-lower-rhine-and-meuse

Peng, F., Prins, M. A., Kasse, C., Cohen, K. M., Van der Putten, N., van der Lubbe, J., Toonen, W. H. J., & van Balen, R. T. (2019). An improved method for paleoflood reconstruction and flooding phase identification, applied to the Meuse River in the Netherlands. Global and Planetary Change177, 213-224. https://doi.org/10.1016/j.gloplacha.2019.04.006

[20] Peng, F., Prins, M. A., Kasse, C., Cohen, K. M., Van der Putten, N., van der Lubbe, J., Toonen, W. H. J., & van Balen, R. T. (2019). An improved method for paleoflood reconstruction and flooding phase identification, applied to the Meuse River in the Netherlands. Global and Planetary Change177, 213-224. https://doi.org/10.1016/j.gloplacha.2019.04.006

[21] Kravanja P, Golob A, Concato M, Leskovar T, Zupanič Pajnič I. Effects of different environmental factors on preservation of DNA in petrous bones: A comparative study of two Slovenian archaeological sites. Forensic Sci Int. 2025 Jun;371:112495. doi: 10.1016/j.forsciint.2025.112495. Epub 2025 May 7. PMID: 40349398 (PubMed) https://pubmed.ncbi.nlm.nih.gov/40349398/

“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. (June 17, 2025). https://www.encyclopedia.com/humanities/encyclopedias-almanacs-transcripts-and-maps/transition-farming-along-lower-rhine-and-meuse

[22] Poetsch M, Markwerth P, Konrad H, Bajanowski T, Helmus J. About the influence of environmental factors on the persistence of DNA – a long-term study. Int J Legal Med. 2022 May;136(3):687-693. doi: 10.1007/s00414-022-02800-6. Epub 2022 Feb 23. PMID: 35195781; PMCID: PMC9005405 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC9005405/

Conor Rossi , Gabriela Ruß-Popa , Valeria Mattiangeli , Fionnuala McDaid , Andrew J. Hare , Hossein Davoudi , Haeedeh Laleh , Zahra Lorzadeh , Roya Khazaeli , Homa Fathi , Matthew D. Teasdale , Abolfazl A’ali , Thomas Stöllner, et al , Exceptional ancient DNA preservation and fibre remains of a Sasanian saltmine sheep mummy in Chehrābād, Iran, Biology Letter, The Royal Society, Jul 2021, 17, 7, https://royalsocietypublishing.org/doi/10.1098/rsbl.2021.0222

[23] Logan Kistler, Roselyn Ware, Oliver Smith, Matthew Collins, Robin G. Allaby, A new model for ancient DNA decay based on paleogenomic meta-analysis, Nucleic Acids Research, Volume 45, Issue 11, 20 June 2017, Pages 6310–6320, https://doi.org/10.1093/nar/gkx361

Conor Rossi , Gabriela Ruß-Popa , Valeria Mattiangeli , Fionnuala McDaid , Andrew J. Hare , Hossein Davoudi , Haeedeh Laleh , Zahra Lorzadeh , Roya Khazaeli , Homa Fathi , Matthew D. Teasdale , Abolfazl A’ali , Thomas Stöllner, et al , Exceptional ancient DNA preservation and fibre remains of a Sasanian saltmine sheep mummy in Chehrābād, Iran, Biology Letter, The Royal Society, Jul 2021, 17, 7, https://royalsocietypublishing.org/doi/10.1098/rsbl.2021.0222

[24] Kravanja P, Golob A, Concato M, Leskovar T, Zupanič Pajnič I. Effects of different environmental factors on preservation of DNA in petrous bones: A comparative study of two Slovenian archaeological sites. Forensic Sci Int. 2025 Jun;371:112495. doi: 10.1016/j.forsciint.2025.112495. Epub 2025 May 7. PMID: 40349398 (PubMed) https://pubmed.ncbi.nlm.nih.gov/40349398/

Poetsch M, Markwerth P, Konrad H, Bajanowski T, Helmus J. About the influence of environmental factors on the persistence of DNA – a long-term study. Int J Legal Med. 2022 May;136(3):687-693. doi: 10.1007/s00414-022-02800-6. Epub 2022 Feb 23. PMID: 35195781; PMCID: PMC9005405 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC9005405/

[24]  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 Iñigo, 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

[25] Ibid

[26] Ibid

[17] Ibid

[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, 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/

Kooijmans, L.P. Louwe, The Rhine/Meuse Delta: Four Studies on its Prehistoric Occupation and Holecene Geology, Lieden University Press, 1974

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)

Kooijmans, L.P. Louwe, The Rhine/Meuse Delta: Four Studies on its Prehistoric Occupation and Holecene Geology, Lieden University Press, 1974

Parasayan O, Laurelut C, Bôle C, Bonnabel L, Corona A, Domenech-Jaulneau C, Paresys C, Richard I, Grange T, Geigl EM. Late Neolithic collective burial reveals admixture dynamics during the third millennium BCE and the shaping of the European genome. Sci Adv. 2024 Jun 21;10(25):eadl2468. doi: 10.1126/sciadv.adl2468. Epub 2024 Jun 19. PMID: 38896620; PMCID: PMC1118650 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC11186501/

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

Pierrik, H.J., E. Stouthamer, K.M. Cohen, Natural levee evolution in the Rhine-Meuse delta, the Netherlands, during the first millennium CE, Geomorphology, 295 (2017) 215-234, https://dspace.library.uu.nl/bitstream/handle/1874/352044/Natural.pdf?sequence=1

Arnoldussen, Stijn, A Living Landscape: Bronze Age Settlement Sites in the Dutch River area (c.2000-80 BC), Sidestone Press, 2008 https://www.sidestone.com/openaccess/9789088900105.pdf

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)

Nienhuis, Piet, H. , Environmental History of the Rhine-Meuse Delta, Springer Science + Business Media B.V., 2008

Amkreutz, L. W. S. W. Persistent traditions: a long-term perspective on communities in the process of Neolithisation in the Lower Rhine Area (5500-2500 cal BC). (Sidestone Press, 2013).

Cromb., P. et al. New evidence on the earliest domesticated animals and possible small-scale husbandry in Atlantic NW Europe. Sci. Rep. 10, 1–15

Brusgaard, N. .. et al. Early animal management in northern Europe: multi-proxy evidence from Swifterbant, the Netherlands. Antiquity 98, 654–671 (2024).

Kooijmans, L. P. L. & Jongste, P. F. B. A neolithic settlement on the Dutch North Sea coast c. 3500 CAL BC. (Analecta Praehistorica Leidensia S., 2006).

Dreshaj, M., Dee, M., Brusgaard, N., Raemaekers, D. & Peeters, H. High resolution Bayesian chronology of the earliest evidence of domesticated animals in the Dutch wetlands (Hardinxveld-Giessendam archaeological sites). PLoS One 18, 1–23 (2023).

[29] See for example

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. A common genetic origin for early farmers from Mediterranean Cardial and Central European LBK cultures. Mol. Biol. Evol. 32, 3132–3142 (2015).

Lazaridis, I. et al. Ancient human genomes suggest three ancestral populations for present-day Europeans. Nature 513, 409–413 (2014). 

Skoglund, P. et al. Origins and genetic legacy of Neolithic farmers and hunter-gatherers in Europe. Science 336, 466–469 (2012).

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.

Miller, Mark, Most European Men are Descended from just Three Bronze Age Warlords, New Study Reveals, 25 may 2015, Ancient Origins, https://www.ancient-origins.net/news-evolution-human-origins/most-european-men-are-descended-just-three-bronze-age-warlords-new-020361

Batini, C., Hallast, P., Zadik, D. et al. Large-scale recent expansion of European patrilineages shown by population resequencing. Nat Commun 6, 7152 (2015). https://doi.org/10.1038/ncomms8152

Abrams, Joel, A handful of Bronze-Age men could have fathered two thirds of Europeans, 21 May 2015, The Conversation, https://theconversation.com/a-handful-of-bronze-age-men-could-have-fathered-two-thirds-of-europeans-42079

[30] Bronze Age deforestation:

Agricultural Expansion: Bronze Age populations in the region expanded upstream along the Rhine and Meuse rivers, leading to extensive forest clearing for farming purposes. The introduction of ploughing techniques also intensified land use and soil disturbance.

Increased Sediment Load: Large-scale deforestation increased sediment runoff into the rivers. By Roman times (2000 years ago), the Rhine and Meuse carried significantly more fine sediment than before, roughly doubling the pre-Bronze Age levels.

Delta Alterations: The increased sediment delivery to the Rhine-Meuse delta caused major changes to the river network. The delta branch network underwent a “complete replacement,” with sediment deposition splitting discharge over more channels and leading to the silting up of older branches. This pulse of sedimentation helped shape the delta as it is known today.

Increased Flood Intensity: The increased sediment load and altered river channels likely led to increased flooding and sedimentation within the delta, potentially ending peat formation in some areas. 

During the Late Bronze Age (1050-800 BC), there is evidence of a decline in settlement sites in the delta, potentially linked to increased flooding and changes in the river system. However, other research suggests that climate change might also have played a role in this decline.

While direct human impact in the delta began later, with activities like peat mining during Roman times and channel embankment in the Middle Ages, the impacts of upstream deforestation in the Bronze Age were already significant. 

Huth, Christoper, Water between two worlds – reflections on the explanatory value of archaeological finds in a Bronze Age river landscape, 276 – 289 in Anne Lehoërff and Marc Talon, Movement, Exchange and Identity in Europe in the 2nd and 1st Millennia BC, Philadelphia: Oxbow s,, 2017, https://www.academia.edu/35831287/Water_between_two_worlds_reflections_on_the_explanatory_value_of_archaeological_finds_in_a_Bronze_Age_river_landscape

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)

Parasayan O, Laurelut C, Bôle C, Bonnabel L, Corona A, Domenech-Jaulneau C, Paresys C, Richard I, Grange T, Geigl EM. Late Neolithic collective burial reveals admixture dynamics during the third millennium BCE and the shaping of the European genome. Sci Adv. 2024 Jun 21;10(25):eadl2468. doi: 10.1126/sciadv.adl2468. Epub 2024 Jun 19. PMID: 38896620; PMCID: PMC1118650 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC11186501/

van Dinter, M., The Roman Limes in the Netherlands: how a delta landscape determined the location of the military structures, Netherlands Journal of Geosciences, 92-1, 11-13 2013, 11-32

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

Pierrik, H.J., E. Stouthamer, K.M. Cohen, Natural levee evolution in the Rhine-Meuse delta, the Netherlands, during the first millennium CE, Geomorphology, 295 (2017) 215-234, https://dspace.library.uu.nl/bitstream/handle/1874/352044/Natural.pdf?sequence=1

Arnoldussen, Stijn, A Living Landscape: Bronze Age Settlement Sites in the Dutch River area (c.2000-80 BC), Sidestone Press, 2008 https://www.sidestone.com/openaccess/9789088900105.pdf

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)

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

[31] van Dinter, M., The Roman Limes in the Netherlands: how a delta landscape determined the location of the military structures, Netherlands Journal of Geosciences, 92-1, 11-13 2013, 11-32

van dinter, Marieke, Living along the Limes Landscape and settlement in the Lower Rhine Delta during Roman and Early Medieval times, PhD Thesis, Utrecht, 2017, https://www.academia.edu/35123998/Living_along_the_Limes_Landscape_and_settlement_in_the_Lower_Rhine_Delta_during_Roman_and_Early_Medieval_times

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

Pierik, H.J., van Lanen, R.J., Roman and early-medieval habitation patterns in a delta landscape: The link between settlement elevation and landscape dynamics, Quaternary International (2017), http://dx.doi.org/10.1016/j.quaint.2017.03.010 

[32] van dinter, Marieke, Living along the Limes Landscape and settlement in the Lower Rhine Delta during Roman and Early Medieval times, PhD Thesis, Utrecht, 2017, https://www.academia.edu/35123998/Living_along_the_Limes_Landscape_and_settlement_in_the_Lower_Rhine_Delta_during_Roman_and_Early_Medieval_times

Pierik, H.J., van Lanen, R.J., Roman and early-medieval habitation patterns in a delta landscape: The link between settlement elevation and landscape dynamics, Quaternary International (2017), http://dx.doi.org/10.1016/j.quaint.2017.03.010 

van Lanen RJ, de Kleijn MTM, Gouw-Bouman MTIJ, Pierik HJ. Exploring Roman and early-medieval habitation of the Rhine–Meuse delta: modelling large-scale demographic changes and corresponding land-use impact. Netherlands Journal of Geosciences. 2018;97(1-2):45-68. doi:10.1017/njg.2018.3, https://www.cambridge.org/core/journals/netherlands-journal-of-geosciences/article/exploring-roman-and-earlymedieval-habitation-of-the-rhinemeuse-delta-modelling-largescale-demographic-changes-and-corresponding-landuse-impact/40F68343AEEC8FF41124C5F098069863

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

[33] The Maastricht-Aachen corridor played a significant role in the early medieval period, particularly during the reigns of the Merovingian and Carolingian dynasties, despite the decline of Roman influence. Maastricht owed its existence to the Via Belgica, a major Roman road connecting Boulogne-sur-Mer to Cologne, as it provided a crucial crossing point of the Meuse river. A Late Roman Castellum (fort) and a bridge across the Meuse were key elements of the city’s infrastructure.

The Maastricht-Aachen corridor, building upon its Roman foundations and strategic location, evolved into a dynamic region in the post-Roman era. It served as a vital center of trade, a political hub, and a significant cultural center during the Merovingian and Carolingian periods, solidifying its place in early medieval European history. 

During the Merovingian Period (450-750 AD), Maastricht became a significant center for trade and commerce due to its strategic location at the crossroads of important routes. The city’s location facilitated the minting and circulation of Merovingian coins throughout Europe. Archeological evidence confirms the presence of artisanal activities, such as pottery production, in Maastricht during this time.

During the Carolingian Period, Maastricht, along with Aachen and Liège, became part of the heartland of the Carolingian dynasty. Aachen evolved into a political and administrative center for Charlemagne.

Maastricht, Wikipedia, This page was last edited on 7 June 2025, https://en.wikipedia.org/wiki/Maastricht

Aachen, Wikipedia, This page was last edited on 8 June 2025,, https://en.wikipedia.org/wiki/Aachen

Arnoldussen, Stijn, A Living Landscape: Bronze Age Settlement Sites in the Dutch River area (c.2000-80 BC), Sidestone Press, 2008 https://www.sidestone.com/openaccess/9789088900105.pdf

Pierik, H.J., van Lanen, R.J., Roman and early-medieval habitation patterns in a delta landscape: The link between settlement elevation and landscape dynamics, Quaternary International (2017), http://dx.doi.org/10.1016/j.quaint.2017.03.010 

[34] Mares, Boed, G-M201, 19 Feb 2025, Marres, https://www.marres.nl/EN/G-M201.htm

Francia, Wikipedia, This page was last edited on 13 June 2025, https://en.wikipedia.org/wiki/Francia

Franks, Wikipedia, This page was last edited on 13 June 2025, https://en.wikipedia.org/wiki/Franks

Altena, E., Smeding, R., van der Gaag, K.J. et al. The Dutch Y-chromosomal landscape. Eur J Hum Genet 28, 287–299 (2020). https://doi.org/10.1038/s41431-019-0496-0

van Es, W.A. and W.J.H. Verwers, Early Medieval settlements along the Rhine: precursors and contemporaries of Dorestad, Journal Archaeology in the Low Countries, 2-1 (May 2010) 5-39, https://jalc.nl/cgi/t/text/get-pdfcfad.pdf?c=jalc%3Bidno%3D0201a01

Uehlinger, Urs, Karl M. Wantzen, Rob S.E.W. Leuven, Hartmut Arndt, The Rhine River Basin, https://d-nb.info/1088795714/34

[35] Ardennes and Eifel are mountain ranges in Europe. “Their western starting point roughly begins where the Meuse river crosses the French-Belgian border. They stretch in a northeastern direction, covering eastern Belgium (Wallonia), northern Luxembourg and western Germany as far as the Rhine river between the cities of Bonn and Koblenz, and are bordered by the Moselle river on the south.

Ardennes and Eifel, Wikipedia, This page was last edited on 1 November 2024, https://en.wikipedia.org/wiki/Ardennes_and_Eifel

Understanding the Phylogenetic ‘Gap’ Associated with the Continental Migratory Route – Part Three

This story is a continuation of my focus on the G Haplogroup phylogenetic tree of the Griff(is)(es)(ith) patrilineal line of descent and the migratory route of the Griffis family Y-DNA in the long term genealogical time layer.

This part of the story focuses on possible macro social-cultural and enviromental influences that limited the growth of YDNA subclades in the migratory path for one of the two major migratory gaps in the family genetic patrilineal line.


The 3,450 year Gap between G-PF3345 and G-L497: The common ancestor associated with the haplogroup G-PF3345 was born around 8550 Before Common Era (BCE). The next common ancestor in my YDNA line was associated with the G-L497 haplogroup who was born about 3,500 years later, around 5525 BCE. If we use 30 years as an estimate of a generation, this gap between major YDNA mutations represents about 115 generations. (See illustration one.)

This phylogenetic gap between haplogroup G-PF3345 and G-L497 represents two endpoints in the migratory route that mirrors the ‘continental migratory route‘ of the Early European Farmers from Anatolia to central Europe. (See illustration one.) The migratory northwestern path between the G-PF3345 and G-L497 essentially followed the Danube River which provided a transport route for generations along a river valley that provided fertile, wind-deposited soils that supported intensive cereal cultivation.

Illustration One: The G-PF3345 – G-L497 Phylogenetic Gap

Click for Larger View | Source: Results for G-Y132595 Haplogroup Migration Using FamilyTreeDNA, Globetrekker

Aside from the methodological factors that may explain the lack of identififed haplogroups in each of these gaps, as outlined in part two of this story, the absence of identified haplogroups in each of these phylogenetic gaps could also be attributed to cultural, social, demographic and environmental influences and factors. 

The Migratory Path of the Phylogenetic Gap Between Haplogroups G-PF3345 and G-L497

Haplogroup G-PF3345 (also known as G2a2b2a1a1) is a significant Y-chromosome haplogroup or YDNA ancestor that represents a majority of haplogroup G descendants in modern Europe. Its origins can be traced to the Neolithic and Chalcolithic periods. [1] . While many ancient G subclades disappeared, G-PF3345 became a dominant surviving lineage in European populations. The common ancestor associated with the defining G-PF3345 Y-DNA SNP mutations lived in the Mesolithic Anatolian area. [2]

The G-PF3345 haplogroup spread across Europe through both Continental and Mediterranean routes during the Neolithic expansion, as evidenced by ancient DNA and modern distribution patterns. G-PF3345 descendants utilized both routes, with the continental Danubian path shaping Central Europe and the Mediterranean coastal path influencing southern Europe. (See illustration two.)

Illustration Two: Migratory Routes of Descendants of G-PF3345

Click for Larger View | Source: Original background map is from Roke,Blank political map of europe, including north africa and western asia, Jun 2006, Wikimedia

The Danubian/Continental Route was associated with a number of Neolithic cultures as the G haplogroup made their way along the Danube River. At the ‘tailend’ of this phylogenetic gap, the Linear Pottery (LBK) culture had a notable presence. This path followed the contours of Danube River and its tributaries into Central Europe (~5,500–4,500 BCE). Farmers settled in regions like Germany, Hungary, and Romania, leaving a genetic legacy in modern populations of Austria, Switzerland, and the Czech Republic. [3]

G2a farmers from the Thessalian Neolithic quickly expanded across the Balkans and the Danubian basin, reaching Serbia, Hungary and Romania by 5800 BCE, Germany by 5500 BCE, and Belgium and northern France by 5200 BCE. Ancient skeletons from the Starčevo–Kőrös–Criș culture (6000-4500 BCE) in Hungary and Croatia, and the Linear Pottery culture (5500-4500 BCE) in Hungary and Germany, all confirmed that G2a (both G2a2a and G2a2b) remained the principal paternal lineage even after farmers intermingled with indigenous populations as they advanced.[4]

The Mediterranean/Coastal Route was linked to the Cardium Pottery culture, this coastal migration moved westward via the Mediterranean (~5,000 BCE), colonizing Italy, southern France, Iberia, and islands like Sardinia and Corsica.

These people crossed the Aegean by boat and colonized the Italian peninsula, the Illyrian coast, southern France and Iberia, where they established the Cardium Pottery culture (5000-1500 BCE). Once again, ancient DNA yielded a majority of G2a samples in the Cardium Pottery culture, with G2a frequencies above 80% (against 50% in Central and Southeast Europe).[5]

Illustration four provides a map of the Near East and Neolithic Europe. The colored areas represent different Neolithic cultures. The black solid lines represent the areas where estimated cultural and chronological distinctions occurred based on archealogical research. The dotted lines encircle the areas associated with each of the two migrator routes. [6]

Illustration Four: European Neolithic Cultures Associated with Continental and Mediterranean Migration Routes

Click for Larger View | Source: A modified version of a map originally found in J. Guilaine, Propagation chronologique, à travers l’Europe, de l’économie néolithique, portée par un certain nombre de cultures “primaires” ou “secondaires” schématiquement cartographiées. Le modèle arythmique.https://traces.univ-tlse2.fr/accueil/equipes-et-ateliers/prbm-prehistoire-recente-du-bassin-mediterraneen/derniers-chasseurs-collecteurs-premiers-agro-pasteurs-transitions-holocenes-en-mediterranee-occidentale

The continental migratory route started northward from Western Anatolia through Greece, and the Southern Balkans (Thrace and Macedonia). [7] The route from Anatolia may have been through settlements of cultures like Karanovo, Hamangia, and Vinčaof, what later became the communities of the LBK (Linear Pottery Culture). The migratory route continued along the Danube River. [8]

The Neolithic Revolution began in the Levant and Anatolia, where domestication of crops like wheat, barley, and legumes, alongside animals such as sheep and goats, laid the foundation for sedentary lifestyles[9] By 7000 BCE, these practices spread northwestward into southeastern Europe, marking the start of the Continental Route, establishing agro-pastoral communities that later influenced the LBK. [10]

The Anatolian farmers’ migration expansion on the continental route was estimated to be  ~0.9 km/year through the Balkans and Central Europe, moving ~50 km per generation. This was slower than the expansion rate associated with the mediterranean route. The continental rate reflects shorter dispersal distances and gradual inland expansion.

A review of various studies on Neolithic expansion indicate the migration of the descendants of the Neolithic Farmers expanded across Europe at an average rate of 1 to 1.3 kilometers per year. This expansion led them to reach the Rhine Valley by around 5300 BCE. [11]

The Danube River is approximately 2,850 kilometers (1,770 miles) long and is part of an extensive water shed (see illustration five). [12] This extensive watershed was the foundation for the Neolithic migration of the G haplogroup into central Europe (see illustration five).

If the estimated expansion rate for the G haplogroup along the Danube River took about 1 to 1.3 km/year, it is conceivable that the it took about 2,565 to 3,700 years to travel the course of the river through successive settlements. The 3,450 year phylogenetic gap coincides with this estimate range of migration expansion.

Illustration Five: The Danube and its Tributaries

Click for Larger View | Source:Danube River Map, Atlas, https://atlas.co/explore/rivers/danube-river/

The second migratory route, the Southern (Maritime) Route, migrated westward along the coastlines of the Adriatic and Mediterranean Seas. They likely used boats and followed the coastlines, establishing settlements along the way. They reached the Paris Basin by around 5000 BCE.  According to some studies, the coastal spread was nearly twice as fast (~1.6 km/year), with farmers traveling ~70 km per generation compared to ~50 km per generation for the contnental route.  In the West Mediterranean, rates exceeded 5 km/year, driven by long-distance maritime voyaging (300–450 km per generation). [13]

The inland route’s slower pace stemmed from overland migration with shorter generational movements, while coastal farmers exploited maritime technology for longer jumps. Coastal dispersal involved leapfrog movements (cabotage), allowing farmers to bypass geographical barriers and establish new settlements rapidly. Simulations show sea travel was essential to explain the West Mediterranean’s rapid spread. [14]

Radiocarbon dates indicate the Neolithic farmers reached Portugal (~2,500 km from the Near East) in ~300 years via the Mediterranean, contrasting with slower continental advances. [15]

The Mediterranean route’s faster spread was enabled by maritime technology and longer dispersal jumps, while the continental route’s slower, steadier progress reflected land-based migration patterns. Both pathways involved similar rates of cultural exchange with hunter-gather haplogroups that resided in Europe, but geographical and behavioral factors shaped distinct genetic and demographic outcomes.

The Mitochondrial haplogroup K (common in early farmers) shows a steeper genetic cline along the continental route compared to the Mediterranean, reflecting differences in dispersal distances. [16] Both routes saw about four percent of farmers interbreeding or acculturating hunter gathering groups per generation. However, the Mediterranean route’s longer coastline led to more cumulative genetic admixture events, resulting in higher hunter gatherer ancestry in regions like Iberia. [17]

Social, Cultural, Demographic and Environmental Explanations of the Gap Between G-PF3345 and G-L497

During this temporal phylogentic gap, Europe experienced significant enviromental and demographic shifts that shaped Y-chromosome diversity in Europe. Between 8000 BCE and 5000 BCE, Europe also underwent transformative social and cultural shifts that profoundly shaped Y-chromosome diversity and subclade formation in YDNA haplogroups.

The lack of subclades in this phylogenetic gap may be attributed to a variety of factors. These factors may have had an independent impact on the G haplogroup at various stages in time. The lack of subclade development was driven by the Neolithic transition to agriculture, the migratory effects on the G haplogroup, the rise of patrilineal social structures, population-demographic movements and climatic changes. The earlier phase of this gap may have involved different mechanisms than those of the later, more socially stratified Neolithic societies.

  • Impact of Initial Early Neolithic Dispersal of Farming Communities (9000-7000 BCE)

The early portions of the phylogenetic time gap likely represents a series of genetic founder effects, bottlenecks, and localized extinctions that occurred during the critical early expansion phase of agriculture in the Anatolia, Aegean Greece and Thrace area before the development of the more complex social hierarchies.

The transition from the hunting and food gathering stage of the Palaeolithic and Mesolithic eras to the agriculutral stages of the Neolithic Period was marked by the organization of settlements with a permanent character and the systematic practice of farming and stock-rearing. The Neolithization [18] of western Anatolia and southeastern Europe was a dynamic and multifaceted process, with overlapping and simultaneous waves of expansion over more than 1,000 years.

The transmission of a Neolithic way of life cannot be reduced to the casual introduction of objects, animals and plants, and to the passive acceptance of these new elements by pre-existing populations. It is an active process, which requires 1. learning new techniques and skills and 2. conforming to a Neolithic mode of thought. Alan Barnard suggests that a foraging lifestyle is, by nature, resilient and resistant to the Neolithic, because foraging ideology differs in almost every aspect to that of farming populations; examples include consumption and saving, decision-making and political hierarchy, universal kinship and degree of kin.

Repeated interactions and exchanges between foraging and farming groups operating as two independent units during a phase of availability, leads to a rapid substitution of resources followed by a phase of consolidation. [19]

An article by Maxime Brami and Volker Heyd, [20] indicates that the movement of early farming groups in the Anatolia region was not a simple, linear process but involved complex interactions, including maritime pioneer colonization and subsequent adoption of agriculture by indigenous populations in the Balkans and Greece. The article revisits and updates the influential hypothesis proposed by James Mellaart [21] which suggested that the earliest Neolithic communities in Greece and the Balkans shared common origins with those in Western Anatolia.

Brami and Heyd argue for a more nuanced view of the Neolithic expansion, emphasizing regional diversity and the importance of Western Anatolia as a dynamic frontier rather than a mere corridor for migration. The spread of farming included the transfer of both material culture and agricultural practices, such as animal husbandry and early dairy technology. The demographic expansion of farming populations from Anatolia into Europe is now understood to have occurred in stages, with Western Anatolia serving as a pivotal intermediary zone.

Small founder groups would establish settlements in new territories. These small pioneer groups carried limited genetic diversity from the outset. Only certain male lineages would be represented in each pioneering settlement. The early phase of the Neolithic transition would have created numerous opportunities for genetic lineage elimination. The small size of pioneering farming communities, coupled with their vulnerability to environmental stresses and potential conflicts, created conditions where entire Y-chromosome lineages could disappear without leaving archaeological traces.

In a similar view, an article by Marek Zvelebil provides a comprehensive review and critical evaluation of the major theories and evidence regarding the transition to agriculture and the emergence of Neolithic societies in Europe. The article synthesizes archaeological, genetic, and social perspectives to argue for a nuanced, regionally variable understanding of how farming spread and Neolithic societies formed. [21a]

Zvelebil emphasizes that the transition to farming in Europe cannot be explained solely by migration (demic diffusion) or by local adoption (indigenist model). Instead, he advocates for a more sophisticated, integrationist view that considers both movement and contact between populations. The process was embedded in pre-existing social and historical contexts, including established networks of contact and the intergenerational transmission of knowledge.

The spread of agriculture and Neolithic society was not uniform across Europe. Instead, it was shaped by local conditions, the history of contact with farming communities from the Near East (Levant and Anatolia), and the agency of both indigenous hunter-gatherers and incoming farmers. The structure (social networks, traditions) and agency (individual and group actions) both played crucial roles in how farming was adopted and adapted.

Zvelebil introduces the concept of “agricultural frontier zones,” regions where foragers and farmers coexisted and interacted over extended periods. These zones were sites of cultural exchange, cooperation, and sometimes conflict, leading to the gradual adoption of farming practices by local populations.

  • The Effects of Boom-and-Bust Population Dynamics at the tail end of this gap

A study by Stephen Shennan and other research colleagues, challenges the assumptions of steady population growth during the Neolithic, highlighting the vulnerability of early farming societies to over-exploitation of local environments. The patterns align with broader Neolithic demographic transitions, where initial agricultural productivity gains were offset by long-term ecological or social strain. The study found no correlation between these demographic fluctuations and known climate events, suggesting endogenous causes like unsustainable resource use, soil depletion, or disease. [22]

The introduction of agriculture in Europe around 8,500 years ago ~6550 BCE, initially led to population booms as farming spread westward and northward. However, regional populations subsequently experienced collapses of 30–60 percent, comparable to the demographic impact of the Black Death, an epidemic that peaked in Europe between 1348 and 1350 CE. [23]

These declines occurred in two distinct waves: the first in Central Europe (~7,500 years ago or ~ 5500 BCE) and the second in Northwest Europe (~6,000 years ago or ~4050 BCE). The first wave of population contraction is around the middle to tail end of the phylogenetic gap that is currently being discussed.

Shennan and his colleagues’ research challenges assumptions of steady population growth during the Neolithic, highlighting the vulnerability of early farming societies to overexploitation of local environments. [24]

  • Environmental Effects, Demographic Bottlenecks and Founder Effects

The ‘8.2 kiloyear event’, a rapid and significant global cooling episode, occurred approximately around 6,200 BC. It lasted between 150 and 400 years. Different areas were affected at different times and in different ways, some areas became cooler and drier, some cooler and wetter. 

The 8.2 kiloyear (ky) event, played a significant role in shaping Early Neolithic population dynamics in the Central Balkans, as outlined by Porčić and his colleagues. The event likely triggered northward migration of farming communities from the southern Balkans/Anatolia into the Central Balkans around 6250 BC, as they sought more favorable conditions amid climatic stress. [25] The Central Balkins is an area just north of Anatolia where haplogroup G-PF33445 orignated.

This northward migration of Anatolian farmers contributed to the first population surge (~6250–6000 BC), which combined high fertility with incoming groups. The 8.2 ky event’s timing aligns with this phase, suggesting environmental pressures accelerated demographic shifts. A population decrease followed around 6000 BC, reaching a low by 5800 BC. A second growth phase (5800–5600 BCE) emerged, attributed primarily to high fertility, before another decline after 5600 BCE.

In Europe, the event had notable ecological and population impacts, such as forest shifts and reductions in Mesolithic populations. The cooling and drying led to significant ecological shifts. Deciduous forests receded in favor of boreal forests in the Alps, while drought conditions in Iberia promoted fire-resistant vegetation. In northeastern Greece, average winter temperatures dropped by over 4°C, likely due to increased influence of the Siberian High. These changes are corroborated by multi-proxy records from caves and sediment cores across southwestern Europe. The cooling and drying led to significant ecological shifts. Deciduous forests receded in favor of boreal forests in the Alps, while drought conditions in Iberia promoted fire-resistant vegetation[26]

Archaeological records in the Balkans and the Aegean suggest that the Neolithic advance of farming paused for a couple of centuries, possibly because cooler and drier conditions reduced crop yields and slowed population growth. This pause may have allowed plants introduced from the Fertile Crescent to acclimatize to new conditions before farming resumed. [27]

The research by Porčić and his associates supports the Neolithic Demographic Transition (NDT) theory, linking population growth to agricultural adoption and sedentism. The Central Balkan trends mirror demographic shifts in western and central Europe, reinforcing the role of demography in Neolithic expansion. [28]

The Neolithic Demographic Transition (NDT) refers to the period of rapid population growth that occurred after the adoption of agriculture by prehistoric societies. This transition purportedly marked by increased birth rates and stable or slightly decreased death rates, led to a significant increase in population size in many parts of the world. [29]

Haplogroup G, particularly sub-clades like G2a, is strongly associated with Neolithic migrations from Anatolia into Europe. However, its phylogenetic tree shows long, narrow branches during this period, reflecting limited genetic diversity despite population expansion.

This apparent paradox of the G haplogroup exhibiting limited genetic diversity and the NDT theory’s contention of rapid population growth associated with agricultural practices can be explained by several ‘countervailing’ factors:

  • Founder Effects of Agricultural Groups During Migration: Neolithic farmers expanding into Europe likely formed small, pioneering groups carrying subsets of G haplogroup diversity. Repeated founder effects the during northwestward migration could have reduced genetic variation in each new population. For example, similar to my G-PF3455 ancestor, the G2a3a-M406 sub-clade shows distinct expansion timelines in Italy (~8,100 years ago) and Iran (~8,800 years ago), aligning with regional Neolithic settlements. These isolated groups preserved specific lineages rather than diversifying broadly. [30]
  • Patrilocal Social Structures: As discussed below, genetic evidence suggests patrilocality (males remaining in their birth communities while females marry in) dominated early farming societies. This practice limited male lineage diversity, as Y-chromosomal lineages like G2a were passed down within localized groups.Despite overall population growth, the effective male population size remained small, amplifying genetic drift and reducing haplogroup branching. [31]
  • Rapid Expansion with Limited Subsequent Diversification: The Neolithic expansion occurred quickly along migration routes (e.g., the Mediterranean coast and Danube Valley), favoring the spread of specific G sub-clades like G-P303 and G-L497. Once established, these lineages dominated local populations without significant later divergence. [32]

Population bottleneck events that created major population and YDNA subclade collapses could have happened due to: climate change events like the 8.2 kiloyear event; the movemenet of small groups migrating along the Danube River water shed, pandemic diseases in farming communities; and conflict and displacement along the migration path. [33]

  • Cultural Practices and Patrilineal Kinship

Research conducted by Monika Karmin and colleagues presents several significant findings that may partly explain the phylogenetic gap. A key discovery from the study was the documentation of a strong bottleneck in male Y-chromosome lineages dating to the last 10,000 years, which contrasts with stable female mitochondrial DNA (mtDNA) patterns. The researchers hypothesize that this recent bottleneck was caused by cultural changes that affected the variance of reproductive success among males. [34]

Illustration six graphically points to the dramatic reduction of male effective population size (Ne) between roughly 8000 BCE to 3000 BCE. Two encircled areas in illustration eleven graphically identify the growth differences in each of the YDNA and mtDNA graphs in this time period.

Illustration Six: Bottleneck of Y Chromosome Diversity Coincides with a Global Change in Culture

Click for Larger View | Source: Karmin M, et al, A recent bottleneck of Y chromosome diversity coincides with a global change in culture. Genome Res. 2015 Apr;25(4):459-66,doi: 10.1101/gr.186684.114, PubMed:https://pmc.ncbi.nlm.nih.gov/articles/PMC4381518/

The decline in the male effective population size (Ne) during the Neolithic period was approximately one-twentieth of its original level in various regions of the world. In the same study, mitochondrial sequences indicated a continual increase in population size from the Neolithic to the present, suggesting extreme divergences between the demographic size of male and female populations in the bottleneck period. [35]

The authors of the study suggest the low estimates of male Ne might be explained by culturally driven sex-specific changes in variance in offspring number rather than natural selectionThis male-specific decline is attributed to cultural shifts tied to the Neolithic transition to agro-pastoralism, which altered male reproductive variance. This bottleneck coincided with global shifts to agriculture in the Near East and Europe between ~10,500 and 6,500 before present (BP) or between ~ 8550 BCE and 4550 BCE. This coincides with the phylogenetic gap between G-PF3345 and G-L497.

This ‘male altered reproductive variance’ arose through two key mechanisms:

1. Patrilineal Social Systems: The rise of patrilocality (males remaining in birthplace) and patrilineal inheritance (father-to-son wealth/status transmission) concentrated reproductive success among high-status males. This reduced effective male population size while female migration maintained mitochondrial diversity. [36] Ancient DNA from Neolithic/Bronze Age sites shows higher male relatedness (shared Y haplogroups) compared to female diversity, consistent with patrilocal practices. [37]

2. Clan Segmentation and Social Stratification: Patrilineal clans splitting into subclans amplified lineage-specific expansions. Social hierarchies allowed dominant lineages to proliferate, while others dwindled. Models show this process alone—without warfare—could explain the genetic bottleneck. [38]

While initial theories proposed intergroup warfare as the driver of the decline in the male effective population size [39] , subsequent research demonstrated that peaceful patrilineal systems could replicate the bottleneck through:

  • Cultural hitchhiking: Y-chromosome lineages became linked to clan identity and success.
  • Non-random lineage loss: Clan expansions/extinctions preferentially affected male genetic diversity. [40]

According to the research arguments, this bottleneck coincided with global shifts to agriculture (Near East/Europe: ~10,500–6,500 BP; East Africa/Arabia: ~7,000–5,000 BP), where wealth accumulation incentivized patrilineal inheritance systems. The delayed genetic signal (3,000–5,000 BP) reflects the time required for these social structures to reshape Y-chromosome diversity.

A Look at Anthropological Evidence for Cultural Explanations of the G-PF3345 and G-L497 Gap

Anthropological evidence suggests that Neolithic societies may have practiced patrilocality, where male descendants remained in ancestral territories. This type of social structural practice could suppress lineage diversification by limiting male mobility and reproductive variance. However, it is not a clear-cut case of causation between patrilocality, the increased stratification of wealth and clan segmentation with the reduction of haplgroup subclades.

While specific cultures from western Turkey between 11,000 and 8,000 BCE are not as well documented as in later periods, the region was home to early human groups transitioning from mobile hunter-gatherers to settled, proto-agricultural communities. This period laid the foundation for the more complex Neolithic cultures that would soon flourish across Anatolia. the Aegean and Baltic areas.

Between 9000 BCE and 5000 BCE, which is roughly the time span of the phylogenetic gap, the Danube River region was a cradle of a number of significant prehistoric cultures, transitioning from Mesolithic hunter-gatherers to early Neolithic farming societies. Based on archaeological evidence and scholarly consensus. Below is an overview in illustration seven of the main cultures that existed along the Danube and in the Anatolian and Baltic areas during this period. Depending on the exact and rate of migration, descendants of G-PF3345 could have been part of any of these cultures.

Illustration Seven: Neolithic Cultures that May have been Associated with Griff(is)(es)(ith) ancestors between the Phylogenetic Gap

The ancestor associated with the G-PF3345 haplogroup was born possibly in Western Anatolia during or after the period when human communities transitioned from hunter-gatherer lifestyles to settled agricultural societies. This time frame encompasses the late Epipaleolithic/Mesolithic, Neolithic, and early Chalcolithic periods, representing one of the most transformative eras in human prehistory. Archaeological evidence reveals a diverse cultural landscape with regional variations, technological innovations, and evolving settlement patterns.

The Aceramic Neolithic tradition extended from approximately 8500-5500 BCE across parts of the Middle East, including portions of western Turkey. This period is characterized by permanent settlements, the beginnings of agriculture and animal domestication, but before the widespread adoption of pottery technology. [41]

During this early phase, the spread of Neolithic lifeways appears to have paused in central Anatolia for over a thousand years, only moving westward toward the Aegean after about 7000 BCE. This suggests a period of cultural adaptation and development before further expansion. [42]

The spread of Neolithic farming in the area where the G-PF3345 ancestor lived in Western Anatolia was not a single, rapid event but occurred in at least two definable waves. The first stage was characterized by sparse and sporadic evidence, lasting until around 6500–6400 BC. The second stage was marked by a more substantial and rapid expansion, especially evident in the lakes district and inner western Anatolia. This movement brought with it a “package” of domesticated plants and animals, ground stone artifacts, and building techniques, but left behind most cult and prestige objects, suggesting the migrants were primarily simple farmers or herdsmen. [43]

One of the earliest Neolithic settlements in western Turkey is the Bilecik-Bahçelievler site in northwestern Anatolia, inhabited between approximately 7100 BCE and 6000 BCE. According to radiocarbon dating evidence, this settlement provides some of the earliest results of the Neolithic period in Western Anatolia and offers important information about the beginning and development of the Fikirtepe Culture, which would later become a dominant cultural tradition in the region. [44]

The spread of fully developed Neolithic cultures into western Anatolia became more pronounced after 6500 BCE, with several distinct cultural traditions emerging across the region. The archaeological evidence reveals complex patterns of cultural interaction and regional differentiation across western Turkey during this period. The spread of the Hacılar culture in the south of Western Anatolia and the spread of the Fikirtepe culture in the north is clearly evident. [45]

The Fikirtepe culture represents one of the most significant Neolithic cultural traditions in northwestern Anatolia. Archaeological research has securely placed this culture between 6450 and 6100 BCE. Subsistence was primarily based on farming, with sheep and goat being more common than cattle, and relatively little evidence of hunting compared to earlier periods and other regions. [46]

Illustration Eight: Core Areas of Neolitic Formation in Anatolia Region

Click for Larger View | Source: Maxime Brami and Volker Heyd, Fig. 16a. Expansion of the DFBW horizon to the region of Marmara leading to the emergence of the ‚Archiac Fikirtepe tradition, The origins of Europe’s first farmers: The role of Hacılar and Western Anatolia, fifty years on, Oct 2011, Praehistorische Zeitschrift 86(2):Page 188, DOI: 10.1515/pz.2011.011

The archaeological sites in illustration eight correspond with the estimated location of the G-PF3455 haplogroup as well as other G haplogroups identified as part of the Griff(is)(es)(ith) migratory path in the Neolithic era. (see illustration nine.)

Illustration Nine: Location of Early G Haplogroups in Western Asia

Click for Larger View | Source: Migratory Path of G Haplogroup Using Terminal Haplogroup G-Y132505 Rendered with Globe Trekker, FamilyTreeDNA, 12 February 2025

A study by Rosenberg and Rocek provides an analysis of Aceramic Neolithic societies in southwestern Asia and challenges traditional assumptions about early social complexity during this time period. Their work emphasizes a complex evolution of socio-political structures, arguing against simplistic models of progression from egalitarian to social hierarchical systems. ‘Heterarchical systems’ prevailed, with multiple overlapping forms of authority (ritual, economic, kinship) rather than centralized hierarchies. Evidence from mega-sites like Göbekli Tepe and Çatalhöyük shows community-based groups coexisting with household-based social structures. [47]

The authors propose that early Neolithic societies developed context-specific solutions to social coordination challenges, blending egalitarian ideals with situational hierarchies. This complexity laid groundwork for later institutionalized inequalities while resisting straightforward categorization into “simple” or “complex” societal types.

Another article by Catherine Twiss and colleagues investigates the nature and extent of social and economic inequality at the Neolithic site of Çatalhöyük in central Anatolia. The study uses a comprehensive, multi-dimensional approach, analyzing a wide range of archaeological data to assess patterns of differentiation among the site’s inhabitants. [48]

The study acknowledges that while some differentiation existed, particularly in symbolic or “prestige” domains, these differences did not amount to entrenched class structures. Çatalhöyük’s society is best characterized as having a “dispersed overlapping mosaic of relationships,” with mechanisms likely in place to suppress or limit the emergence of pronounced hierarchies. Whether the nascent agrarian cultures and social structures in the Aceramic Neolithic societies had an impact on the structure of G haplogroup is not certain.

The Neolithic revolution, marked by the adoption of agriculture and settled life, began spreading into the Danube basin from the southeast around 6200–5500 BCE. The Starčevo culture (ca. 6200–5500 BCE) is recognized as the earliest Neolithic culture in the central Danubian region, covering parts of present-day Serbia, Hungary, and Croatia. [49]

In the lower Danube region (modern Bulgaria), the Karanovo culture (ca. 6200–5500 BCE and onward) was another major early Neolithic society, closely related to the spread of farming and settlement patterns in the Balkans. [50] The Dudeşti and Criș Culture in Romania area are contemporaneous with Starčevo and contributed to the Neolithic mosaic along the Danube, particularly in the lower reaches.

Illustration Ten: Extent of the Starčevo–Kőrös–Criș culture (c. 6200-4500 BCE)

Click for Larger View | Source: Maciaomo Hay,Extent of the Starčevo–Kőrös–Criș culture, (c. 6200-4500 BCE), Starčevo–Kőrös–Criș culture, Eupedia,https://www.eupedia.com/genetics/starcevo_culture.shtml

After 5000 BCE, the region saw the rise of even more complex societies, such as the GumelniţaCucuteni-Tripolye, and Varna cultures, which are famous for their metallurgy, large settlements, and elaborate burials (see table one and illustration eleven below). [51]

Table One: Neolithic Cultures in the Danube River Valley

PeriodCulture(s)Location/ExtentKey Features
8000–6000 BCEMesolithic groupsDanube valley (Austria, Hungary, etc.)Hunter-gatherers, flint tools, fishing, cremation
6200–5500 BCEStarčevo, KaranovoBalkans, Central DanubeFirst farmers, pottery, animal domestication
5500–5000 BCELBK (Danubian)Central/Eastern Europe, Danube basinLinear pottery, longhouses, intensive agriculture
5700–4500 BCEVinčaMiddle Danube (Serbia, Balkans)Large settlements, copper use, early writing
ca. 5500 BCEDudeşti, CrișLower Danube (Romania)Early Neolithic farming, regional pottery styles

Illustration Ten: Chalcolithic Cultures of Southeastern Europe with Major Archaeological Sites

Click for Larger View | Source: Modified version of map found at Caliniuc, Eneolithic cultures of Southeastern Europe,13 August 2010, Wikimedia Commons,https://commons.wikimedia.org/wiki/File:SEE-Eneolithic-cultures-Cucuteni.jpg

The Cucuteni-Trypillia culture (CTC), linked to G-PF3345, exhibited large, stable settlements that may have reinforced such practices, reducing opportunities for new subclades to emerge and persist. [52]

While the CTC existed during the Y-chromosome bottleneck period, its genetic record does not strongly align with the social or demographic mechanisms driving that bottleneck (e.g., patrilineal kinship, steppe male expansions). CTC settlements show no clear evidence of patrilineal kinship systems—a key driver of the Y-chromosome bottleneck. The culture’s large, planned settlements (e.g., 15,000-resident “mega-sites”) imply collective social organization rather than lineage-based hierarchies. [53]

The culture’s Y-DNA diversity—particularly the persistence of G2a and I2a—suggests it was peripheral to the male-lineage homogenization occurring in steppe-influenced societies. The bottleneck’s primary documentation remains associated with later Indo-European cultures (Steppe), not the CTC. [54]

Researchers have proposed that the formation of patrilineal kin groups and competition between these groups led to a significant reduction in Y-chromosomal diversity through a process called ‘cultural hitchhiking’. In segmentary patrilineal systems, closely related males cluster together in descent groups. Combined with variance in reproductive success between groups, this can substantially reduce Y-chromosome diversity without requiring violence between groups. Patrilineal systems reduced Y diversity within 70–200 generations of their adoption, aligning with Neolithic social changes. [55]

In some societies, particularly after the development of agriculture and herding, a small number of males may have had disproportionate reproductive success, limiting the diversity of YDNA lineages. This social structure could have allowed certain lineages to dominate, potentially eliminating alternative branches that might have otherwise developed into intermediate subclades.

DNA data suggests dates and places for early PF3345 that correspond with the Neolithic Cucuteni Tripolye culture of Romania and Moldova, with two successful branches developing during the 4th millennium BC -An Eastward expansion (branches of U1) towards the Caucasus and a more general Westward migration towards Southern Germany.” [56]

The roots of Cucuteni–Trypillia culture can be found in the Starčevo–Körös–Criș and Vinča cultures of the 6th to 5th millennia, with additional influence from the Bug–Dniester culture (6500–5000 BC). During the early period of its existence (in the fifth millennium BC), the Cucuteni–Trypillia culture was also influenced by the Linear Pottery culture from the north, and by the Boian culture from the south.[57]

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. [58] 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. [59]

The LBK shows strong archaeological evidence for patrilocality (males remaining in their birth communities) and patrilineal wealth inheritance, supported by isotopic, genetic, and mortuary analyses. Strontium isotope studies of LBK burials reveal significantly less geographic variance among males compared to females, indicating males typically resided in their birthplace while females migrated from other communities. This pattern aligns with a patrilocal kinship system. [60] Grave goods such as polished adzes, flint tools, and Spondylus shell ornaments disproportionately accompanied male burials, reflecting intergenerational wealth transmission. These items symbolized agricultural authority and social status. [61] Genetic analyses of LBK populations show Y-chromosome haplogroups (e.g., G2a, I2) passed through male lines, consistent with patrilineal descent. Conversely, mitochondrial DNA diversity suggests female exogamy. [62]

LBK longhouses likely housed multigenerational male kin groups, with land and resources controlled patriarchally[63] The standardization of house architecture and tool traditions over centuries implies stable male-dominated inheritance practices. [64] Empty grave plots in LBK cemeteries may represent symbolic claims to lineage-based land rights, further reinforcing hereditary wealth structures.[65] This evidence collectively underscores a society where male lineage dictated resource access and social standing, with women integrating into new communities through marriage.

Conclusion and the Next Phylogenetic Gap

The Griff(is)(es)(ith) genetic ancestral paternal line was associated with Neolithic migrations from Anatolia into Europe. However, its phylogenetic tree shows 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.

This limited genetic diversity and lack of documented haplgroups along this migratory path can be explained by several envronmental, demographic, social and cultural ‘countervailing’ factors.

The next and final part of this story on the phyogentic structure of the Griff(is)(es)(ith) genetic ancestral paternal line focuses on the second gap between between G-FGC7516 and G-Z6748.

Source:

Feature Banner: The banner at the top of the story features a map of the phylogenetic gap discussed in the story. The map was generated by taking snapshops from the FamilyTreeDNA GlobetrekkerTM 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.

[1] The enigma of G-PF3345 (U1, CTS342 and L497), 3 Sep 2018, G2a, YDNA Haplogroups, Population Genetics, Forums, Eupedia, https://www.eupedia.com/forum/threads/the-enigma-of-g-pf3345-u1-cts342-and-l497.37040/

[2] Mares, Boed, G-M201, 19 Feb 2025, Marres, https://www.marres.nl/EN/G-M201.htm

Genetic studies on Croats, Wikipedia, This page was last edited on 10 March 2025, https://en.wikipedia.org/wiki/Genetic_studies_on_Croats

Di Cristofaro J, Mazières S, Tous A, Di Gaetano C, Lin AA, Nebbia P, Piazza A, King RJ, Underhill P, Chiaroni J. Prehistoric migrations through the Mediterranean basin shaped Corsican Y-chromosome diversity. PLoS One. 2018 Aug 1;13(8):e0200641. doi: 10.1371/journal.pone.0200641. PMID: 30067762; PMCID: PMC6070208. (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC6070208/

Your Haplogroup Story: G-PF3345 , FamilyTreeDNA , https://discover.familytreedna.com/y-dna/G-PF3345/story

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

Hay, Maciamo, Linear Pottery (LBK) culture (c. 5600-4250 BCE), Eupedia, https://www.eupedia.com/genetics/linear_pottery_culture.shtml

[3] Fort, J., Pérez-Losada, J. Interbreeding between farmers and hunter-gatherers along the inland and Mediterranean routes of Neolithic spread in Europe. Nat Commun 15, 7032 (2024). https://doi.org/10.1038/s41467-024-51335-4

N. Isern, J. Zilhão, J. Fort, & A.J. Ammerman, Modeling the role of voyaging in the coastal spread of the Early Neolithic in the West Mediterranean, Proc. Natl. Acad. Sci. U.S.A. 114 (5) 897-902, 2017 https://doi.org/10.1073/pnas.1613413114 

[4] Hay, Maciamo, Linear Pottery (LBK) culture (c. 5600-4250 BCE), Eupedia, https://www.eupedia.com/genetics/linear_pottery_culture.shtml

[5] Ibid

[6] The name and definition of the color coded areas in the map are below, with references:

Pre-Pottery Neolithic (PPNB) – Pre-Pottery Neolithic B, Wikipedia, This page was last edited on 26 January 2025, https://en.wikipedia.org/wiki/Pre-Pottery_Neolithic_B

Early Neolithic of Northwestern Anatolia – See for example Karul, Necmi. (2022). The Beginning and the Development of Farming-Based Village Life in Northwestern Anatolia / 2022. 231-246. 10.1017/9781107337640.017 https://www.researchgate.net/publication/360398204_The_Beginning_and_the_Development_of_Farming-Based_Village_Life_in_Northwestern_Anatolia_2022

Monochrome / Proto-Sesklo – Sesklo, Wikipedia, This page was last edited on 22 October 2024, https://en.wikipedia.org/wiki/Sesklo

Karanovo – Karanovo culture, Wikipedia, This page was last edited on 23 March 2025, https://en.wikipedia.org/wiki/Karanovo_culture

Anzabegovo – Marco Porcic, Evaluating Social Complexity and Inequality in the Balkans Between 6500 and 4200 BC, Journal of Archaeological Research 27(3):335-390 DOI:10.1007/s10814-018-9126-6, https://www.researchgate.net/publication/327425832_Evaluating_Social_Complexity_and_Inequality_in_the_Balkans_Between_6500_and_4200_BC

Starčevo – Starčevo culture, Wikipedia, This page was last edited on 29 November 2024, https://en.wikipedia.org/wiki/Starčevo_culture

Bug-Dniester  Bug – Dniester culture, Wikiedia, This page was last edited on 2 October 2024, https://en.wikipedia.org/wiki/Bug–Dniester_culture

Rubané Neolithic – Linear Pottery culture (LBK), Wikipedia, This page was last edited on 3 April 2025, https://en.wikipedia.org/wiki/Linear_Pottery_culture

Adriatic Impressa- Cardium pottery, Wikipedia, This page was last edited on 24 February 2025, This page was last edited on 24 February 2025, https://en.wikipedia.org/wiki/Cardium_pottery

Cardial and Derivatives – The term “Cardial” refers to a specific pottery style and associated culture of the Early Neolithic period in Southern Europe, particularly along the Mediterranean coast. Cardial pottery is characterized by distinctive impressions, often made using the edges of Cardium shells, and is a key marker of the Cardial culture. Derivatives of Cardial, such as Epicardial, represent later developments or variations within the Cardial tradition. See Cardium pottery, Wikipedia, This page was last edited on 24 February 2025, This page was last edited on 24 February 2025, https://en.wikipedia.org/wiki/Cardium_pottery

Epicardial – Elsa Defranould. The Cardial–Epicardial Early Neolithic of Lower Rhône Valley (South-Eastern France): A Lithic Perspective. Open Archaeology, 2021, 7 (1), pp.939-952. ⟨10.1515/opar-2020-0182⟩. ⟨hal-04913839⟩. https://hal.science/hal-04913839

Hoguette – La Hoguette, Wikipedia, This page was last edited on 15 April 2025, https://en.wikipedia.org/wiki/La_Hoguette

[7] Thrace refers to a historical region in Southeast Europe, now divided between Bulgaria, Greece, and Turkey. Thrace today is divided among three modern nations: southeastern Bulgaria (Northern Thrace), northeastern Greece (Western Thrace), and the European part of Turkey (East Thrace). The historical region of Thrace encompassed these areas and more, extending along the Balkan Peninsula. 

Macedonia refers to both a historical region and a modern country in Southeastern Europe. Historically, it encompasses an area including parts of Greece, Bulgaria, and the modern-day Republic of North Macedonia. The Republic of North Macedonia, formerly known as the Republic of Macedonia, is a landlocked country with a diverse history and culture, bordering Greece, Albania, Bulgaria, Kosovo, and Serbia. 

Macedonia, Wikipedia, This page was last edited on 30 April 2025, https://en.wikipedia.org/wiki/Macedonia

Thracia, Wikipedia, This page was last edited on 28 January 2025, https://en.wikipedia.org/wiki/Thracia

Thrace, Wikipedia, This page was last edited on 24 March 2025, https://en.wikipedia.org/wiki/Thrace

[8] Neolithic Revolution, Wikipedia, This page was last edited on 16 April 2025, https://en.wikipedia.org/wiki/Neolithic_Revolution

[9] Neolithic, Wikipedia, This page was last edited on 26 April 2025, https://en.wikipedia.org/wiki/Neolithic

[10] Szécsényi-Nagy A, Brandt G, Haak W, Keerl V, Jakucs J, Möller-Rieker S, Köhler K, Mende BG, Oross K, Marton T, Osztás A, Kiss V, Fecher M, Pálfi G, Molnár E, Sebők K, Czene A, Paluch T, Šlaus M, Novak M, Pećina-Šlaus N, Ősz B, Voicsek V, Somogyi K, Tóth G, Kromer B, Bánffy E, Alt KW. 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/

Szécsényi-Nagy A, et al, Tracing the genetic origin of Europe’s first farmers reveals insights into their social organization, 22 April 2015, Proceedings of the Royal Society Biological, https://doi.org/10.1098/rspb.2015.0339, https://royalsocietypublishing.org/doi/10.1098/rspb.2015.0339

[11] Fort, J., Pérez-Losada, J. Interbreeding between farmers and hunter-gatherers along the inland and Mediterranean routes of Neolithic spread in Europe. Nat Commun 15, 7032 (2024). https://doi.org/10.1038/s41467-024-51335-4

Pinhasi, Ron, Joaquin Fort, Albert J Ammerman, Tracing the Origin and Spread of Agriculture in Europe, 29 Nov 2005, PLOS, https://doi.org/10.1371/journal.pbio.0030410

Alexandros Tsoupas, Carlos S. Reyna-Blanco, Claudio S. Quilodrán, Jens Blöcher, Maxime Brami, Daniel Wegmann, Joachim Burger, Mathias Currat, Local increases in admixture with hunter-gatherers followed the initial expansion of Neolithic farmers across continental Europe, bioRxiv, 12 Jun 2024, doi: https://doi.org/10.1101/2024.06.10.598301 , https://www.biorxiv.org/content/10.1101/2024.06.10.598301v1.full.pdf

[12] Danube River Map, Atlas, https://atlas.co/explore/rivers/danube-river/

[13] N. Isern, J. Zilhão, J. Fort, & A.J. Ammerman,  Modeling the role of voyaging in the coastal spread of the Early Neolithic in the West Mediterranean, Proc. Natl. Acad. Sci. U.S.A. 114 (5) 897-902, 2017, https://doi.org/10.1073/pnas.1613413114 .

Fort, J., Pérez-Losada, J. Interbreeding between farmers and hunter-gatherers along the inland and Mediterranean routes of Neolithic spread in Europe. Nat Commun 15, 7032 (2024). https://doi.org/10.1038/s41467-024-51335-4

Neolithic Revolution, Wikipedia, This page was last edited on 16 April 2025, https://en.wikipedia.org/wiki/Neolithic_Revolution

Fort, J., Pérez-Losada, J. Interbreeding between farmers and hunter-gatherers along the inland and Mediterranean routes of Neolithic spread in Europe. Nat Commun 15, 7032 (2024). https://doi.org/10.1038/s41467-024-51335-4

Neolithic Revolution, Wikipedia, This page was last edited on 16 April 2025, https://en.wikipedia.org/wiki/Neolithic_Revolution

By comparing Neolithic spread rates in several regions of the world, Joaquim Fort proposed the following ‘ laws of Neolithic Expansion’:

  1. The Neolithic spread inland at a rate of about 1 km/yr, but there was substantial variation (0.44-3.6 km/yr)
  2. When in addition to demic diffusion there is substantial cultural diffusion, Neolithic spread rates are faster. 
  3. Neolithic spread rates over the sea take place at about 10 km/yr. 
  4. Most inland and coastal Neolithic spreads were mainly demic. 
  5. Neolithic spread rates tend to become slower at higher latitudes. 
  6. The Neolithic spreads later and more slowly at higher altitudes above sea level (compared to surrounding regions). A spatial interpolation of early Neolithic dates in Europe has made it possible to map the isochrones every 250 years and this has shown that the Neolithic first surrounded the Alps completely, and only later begun to climb up these mountains 

Fort, Joaquim, Prehistoric spread rates and genetic clines, 6 Apr 2022, Human Population Genetic and Genomics, 2022; 2(2):0003,  https://www.pivotscipub.com/hpgg/2/2/0003

See also:

Aoki, Kenici, Interpreting the demic diffusion of early farming in Europe with a three-population model , 8 Oct 2024 ,Human Population Genetics and Genomics, 2024;4(4):0010,   https://doi.org/10.47248/hpgg2404040010  

Michael Kempf, Solène Denis, Resource dependency and communication networks in Early Neolithic western Europe, Quaternary Environments and Humans,
Volume 2, Issue 5, 2024, 100014, ISSN 2950-2365, https://doi.org/10.1016/j.qeh.2024.100014 .
(https://www.sciencedirect.com/science/article/pii/S2950236524000124 )

Marko Porčića,Tamara Blagojević, Jugoslav Pendić,Sofija Stefanović, The timing and tempo of the Neolithic expansion across the Central Balkans in the light of the new radiocarbon evidence, Journal of Archaeological Science: Reports, Vol 33, Oct 2020, 102528, 1 – 12, https://www.sciencedirect.com/science/article/pii/S2352409X20303199

Peter Rowley-Conwy, Westward Ho! The Spread of Agriculture from Central Europe to the Atlantic, Current Anthropology Volume

[14]  N. Isern, J. Zilhão, J. Fort, & A.J. Ammerman,  Modeling the role of voyaging in the coastal spread of the Early Neolithic in the West Mediterranean, Proc. Natl. Acad. Sci. U.S.A. 114 (5) 897-902, 2017, https://doi.org/10.1073/pnas.1613413114 .

Paschou P, Drineas P, Yannaki E, Razou A, Kanaki K, Tsetsos F, Padmanabhuni SS, Michalodimitrakis M, Renda MC, Pavlovic S, Anagnostopoulos A, Stamatoyannopoulos JA, Kidd KK, Stamatoyannopoulos G. Maritime route of colonization of Europe. Proc Natl Acad Sci U S A. 2014 Jun 24;111(25):9211-6. doi: 10.1073/pnas.1320811111. Epub 2014 Jun 9. PMID: 24927591; PMCID: PMC4078858, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC4078858/

[15] N. Isern, J. Zilhão, J. Fort, & A.J. Ammerman, Modeling the role of voyaging in the coastal spread of the Early Neolithic in the West Mediterranean, Proc. Natl. Acad. Sci. U.S.A. 114 (5) 897-902 2017, ,https://doi.org/10.1073/pnas.1613413114 

Paschou P, et al, Maritime route of colonization of Europe. Proc Natl Acad Sci U S A. 2014 Jun 24;111(25):9211-6. doi: 10.1073/pnas.1320811111. Epub 2014 Jun 9. PMID: 24927591; PMCID: PMC4078858, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC4078858/

[16] Fort, J., Pérez-Losada, J. Interbreeding between farmers and hunter-gatherers along the inland and Mediterranean routes of Neolithic spread in Europe. Nat Commun 15, 7032 (2024). https://doi.org/10.1038/s41467-024-51335-4

[17] Fort, J., Pérez-Losada, J. Interbreeding between farmers and hunter-gatherers along the inland and Mediterranean routes of Neolithic spread in Europe. Nat Commun 15, 7032 (2024). https://doi.org/10.1038/s41467-024-51335-4

Alexandros Tsoupas, Carlos S. Reyna-Blanco, Claudio S. Quilodrán, Jens Blöcher, Maxime Brami, Daniel Wegmann, Joachim Burger, Mathias Currat, Local increases in admixture with hunter-gatherers followed the initial expansion of Neolithic farmers across continental Europe, bioRxiv, 12 Jun 2024, doi: https://doi.org/10.1101/2024.06.10.598301 , https://www.biorxiv.org/content/10.1101/2024.06.10.598301v1.full.pdf

Paschou P, Drineas P, Yannaki E, Razou A, Kanaki K, Tsetsos F, Padmanabhuni SS, Michalodimitrakis M, Renda MC, Pavlovic S, Anagnostopoulos A, Stamatoyannopoulos JA, Kidd KK, Stamatoyannopoulos G. Maritime route of colonization of Europe. Proc Natl Acad Sci U S A. 2014 Jun 24;111(25):9211-6. doi: 10.1073/pnas.1320811111. Epub 2014 Jun 9. PMID: 24927591; PMCID: PMC4078858 (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC4078858/

Szécsényi-Nagy A, et al, Tracing the genetic origin of Europe’s first farmers reveals insights into their social organization, 22 April 2015, Proceedings of the Royal Society Biological, https://doi.org/10.1098/rspb.2015.0339, https://royalsocietypublishing.org/doi/10.1098/rspb.2015.0339

Maïté Rivollat et al. , Ancient genome-wide DNA from France highlights the complexity of interactions between Mesolithic hunter-gatherers and Neolithic farmers.Sci. Adv.6,eaaz5344(2020).DOI:10.1126/sciadv.aaz5344

e 52, Supplement 4, October 2011, S431-451, https://www.journals.uchicago.edu/doi/epdf/10.1086/65836

[18] Neolithization refers to the process of a society transitioning from a hunter-gatherer lifestyle to a settled agricultural one, marking the transition to the Neolithic period. It essentially describes the widespread adoption of farming and the establishment of villages, signifying a fundamental shift in human societies. 

Click for Larger View | Source:Goce Naumov, The Early Neolithic communities in Macedonia, Archeologické rozhledy LXVII–2015, 331-355 , DOI: 10.35686/AR.2015.18, https://www.researchgate.net/publication/292392154

Goce Naumov, The Early Neolithic communities in Macedonia, Archeologické rozhledy LXVII–2015,  331-355 , DOI: 10.35686/AR.2015.18,  https://www.researchgate.net/publication/292392154 

See also Todd Paradine, Haplogroup G in the New Stone Age, April 2021, GM3302, https://sites.google.com/view/gm3302/the-story/neolithic; PDF verson

[19] Maxime Brami and Volker Heyd, The origins of Europe’s first farmers: The role of Hacılar and Western Anatolia, fifty years on, Oct 2011, Praehistorische Zeitschrift 86(2) 193, DOI: 10.1515/pz.2011.011

[20] Maxime Brami and Volker Heyd, The origins of Europe’s first farmers: The role of Hacılar and Western Anatolia, fifty years on, Oct 2011, Praehistorische Zeitschrift 86(2)165-206, DOI: 10.1515/pz.2011.011

See also M. Brami, ‘Aegean’ and ‘Anatolian’ first farmers: ambiguous labelling or research blind spot. In: A. Lahelma, M. Ahola, E. Holmqvist-Sipilä, K. Mannermaa, K. Nordqvist eds., Moving northward: Professor Volker Heyd’s Festschrift as he turns 60 (Archaeological Society of Finland) 209-219 https://www.academia.edu/106735707/_2023_M_Brami_Aegean_and_Anatolian_first_farmers_ambiguous_labelling_or_research_blind_spot_In_A_Lahelma_M_Ahola_E_Holmqvist_Sipilä_K_Mannermaa_K_Nordqvist_eds_Moving_northward_Professor_Volker_Heyds_Festschrift_as_he_turns_60_Archaeological_Society_of_Finland_209_219

[21] Mellaart, James, Excavations at Hacılar: Third Preliminary Report, Anatolian Stud. 10, 1960, 83–104

Mellaart, James, Excavations at Halicar, Edinburgh, Published for British Institute of Archaeology at Ankara. Edinburgh University Press, 1970 https://archive.org/details/excavationsathac0002mell/page/n3/mode/2up

See also Shennan S. The Origins of Agriculture in South-West Asia. In: The First Farmers of Europe: An Evolutionary Perspective. Cambridge World Archaeology. Cambridge University Press; 2018:16-54.

[21a] Zvelebil, Marek, The agricultural transition and the origins of Neolithic society in Europe, Docummenta Praehistorica, 22 Dec 2001, 28, 1-26. https://doi.org/10.4312/dp.28.1

[22] Shennan S, Downey SS, Timpson A, Edinborough K, Colledge S, Kerig T, Manning K, Thomas MG. Regional population collapse followed initial agriculture booms in mid-Holocene Europe. Nat Commun. 2013;4:2486. doi: 10.1038/ncomms3486. PMID: 24084891; PMCID: PMC3806351, (PubMed) https://pubmed.ncbi.nlm.nih.gov/24084891/

Bower, Bruce, Ancient farming populations went boom, then bust, 1 Oct 2013, Science News, https://www.sciencenews.org/article/ancient-farming-populations-went-boom-then-bust

[23] Porčić Marko, Blagojević Tamara, Pendić Jugoslav and Stefanović Sofija, The Neolithic Demographic Transition in the Central Balkans: population dynamics reconstruction based on new radiocarbon evidence Phil. Trans. R. Soc. 30 Nov 2020, B37620190712 http://doi.org/10.1098/rstb.2019.0712

[24] Shennan S, Downey SS, Timpson A, Edinborough K, Colledge S, Kerig T, Manning K, Thomas MG. Regional population collapse followed initial agriculture booms in mid-Holocene Europe. Nat Commun. 2013;4:2486. doi: 10.1038/ncomms3486. PMID: 24084891; PMCID: PMC3806351, (PubMed) https://pubmed.ncbi.nlm.nih.gov/24084891/

Bower, Bruce, Ancient farming populations went boom, then bust, 1 Oct 2013, Science News, https://www.sciencenews.org/article/ancient-farming-populations-went-boom-then-bust

[25] Porčić Marko, et al, The Neolithic Demographic Transition in the Central Balkans: population dynamics reconstruction based on new radiocarbon evidence Phil. Trans. R. Soc. 30 Nov 2020, B37620190712 http://doi.org/10.1098/rstb.2019.0712

[26] 8.2-kiloyear event, Wikipedia, This page was last edited on 28 March 2025, https://en.wikipedia.org/wiki/8.2-kiloyear_event

García-Escárzaga A, Gutiérrez-Zugasti I, Marín-Arroyo AB, Fernandes R, Núñez de la Fuente S, Cuenca-Solana D, Iriarte E, Simões C, Martín-Chivelet J, González-Morales MR, Roberts P. Human forager response to abrupt climate change at 8.2 ka on the Atlantic coast of Europe. Sci Rep. 2022 Apr 20;12(1):6481. doi: 10.1038/s41598-022-10135-w. PMID: 35444222; PMCID: PMC9021199, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC9021199/

King, David T., 8.2 kiloyear event, 2025, EBSCO, https://www.ebsco.com/research-starters/earth-and-atmospheric-sciences/82-kiloyear-event

Dixit, Y., Chua, S., Yan, Y.T. et al. Hydroclimatic impacts of the abrupt cooling event 8200 years ago in the western Indo-Pacific Warm Pool. Commun Earth Environ 5, 690 (2024). https://doi.org/10.1038/s43247-024-01825-6

Hege Kilhavn, Isabelle Couchoud, Russell N. Drysdale, Carlos Rossi, John Hellstrom, Fabien Arnaud, and Henri Wong, The 8.2 ka event in northern Spain: timing, structure and climatic impact from a multi-proxy speleothem record, Clim. Past, 18, 2321–2344, 2022 , https://doi.org/10.5194/cp-18-2321-2022 and https://cp.copernicus.org/articles/18/2321/2022/cp-18-2321-2022.pdf

Nick Nutter , Nick, Climatic Events that Changed the World: The 8.2k yr BP climate event, Last Updated 29 Jan 2024, Nutter’s World, https://nuttersworld.com/climactic-events/8.2k-yr-event/

Kilhavn, H., Couchoud, I., Drysdale, R. N., Rossi, C., Hellstrom, J., Arnaud, F., and Wong, H.: The 8.2 ka event in northern Spain: timing, structure and climatic impact from a multi-proxy speleothem record, Clim. Past, 18, 2321–2344, https://doi.org/10.5194/cp-18-2321-2022, 2022

[27] Porčić Marko, Blagojević Tamara, Pendić Jugoslav and Stefanović Sofija, The Neolithic Demographic Transition in the Central Balkans: population dynamics reconstruction based on new radiocarbon evidence Phil. Trans. R. Soc. 30 Nov 2020, B37620190712 http://doi.org/10.1098/rstb.2019.0712

[28] Marko Porčić ,Tamara Blagojević , Sofija Stefanović Demography of the Early Neolithic Population in Central Balkans: Population Dynamics Reconstruction Using Summed Radiocarbon Probability Distributions , August 10, 2016, PLOS,  https://doi.org/10.1371/journal.pone.0160832

[29] 8.2-kiloyear event, Wikipedia, This page was last edited on 28 March 2025, https://en.wikipedia.org/wiki/8.2-kiloyear_event

García-Escárzaga A, Gutiérrez-Zugasti I, Marín-Arroyo AB, Fernandes R, Núñez de la Fuente S, Cuenca-Solana D, Iriarte E, Simões C, Martín-Chivelet J, González-Morales MR, Roberts P. Human forager response to abrupt climate change at 8.2 ka on the Atlantic coast of Europe. Sci Rep. 2022 Apr 20;12(1):6481. doi: 10.1038/s41598-022-10135-w. PMID: 35444222; PMCID: PMC9021199, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC9021199/

[30] Rootsi, S., Myres, N., Lin, A. et al. Distinguishing the co-ancestries of haplogroup G Y-chromosomes in the populations of Europe and the Caucasus. Eur J Hum Genet 20, 1275–1282 (2012). https://doi.org/10.1038/ejhg.2012.86

Balaresque P, Bowden GR, Adams SM, Leung H-Y, King TE, Rosser ZH, et al. (2010) A Predominantly Neolithic Origin for European Paternal Lineages. PLoS Biol 8(1): e1000285. https://doi.org/10.1371/journal.pbio.1000285

[31] Szécsényi-Nagy A, et al, Tracing the genetic origin of Europe’s first farmers reveals insights into their social organization, 22 April 2015, Proceedings of the Royal Society Biological, https://doi.org/10.1098/rspb.2015.0339, https://royalsocietypublishing.org/doi/10.1098/rspb.2015.0339

[32] Rootsi, S., Myres, N., Lin, A. et al. Distinguishing the co-ancestries of haplogroup G Y-chromosomes in the populations of Europe and the Caucasus. Eur J Hum Genet 20, 1275–1282 (2012). https://doi.org/10.1038/ejhg.2012.86

Szécsényi-Nagy A, et al, Tracing the genetic origin of Europe’s first farmers reveals insights into their social organization, 22 April 2015, Proceedings of the Royal Society Biological, https://doi.org/10.1098/rspb.2015.0339, https://royalsocietypublishing.org/doi/10.1098/rspb.2015.0339

[33] See the following for evidence for disease-driven population collapses:


Rascovan, N., et al. (2019). “Emergence and spread of basal lineages of Yersinia pestis during the Neolithic decline.” Cell, 176(1-2), 295-305. and Rascovan N, Sjögren KG, Kristiansen K, Nielsen R, Willerslev E, Desnues C, Rasmussen S. Emergence and Spread of Basal Lineages of Yersinia pestis during the Neolithic Decline. Cell. 2019 Jan 10;176(1-2):295-305.e10. doi: 10.1016/j.cell.2018.11.005. Epub 2018 Dec 6. PMID: 30528431. (PubMed) https://pubmed.ncbi.nlm.nih.gov/30528431/ or https://www.cell.com/cell/fulltext/S0092-8674(18)31464-8?_returnURL=https%3A%2F%2Flinkinghub.elsevier.com%2Fretrieve%2Fpii%2FS0092867418314648%3Fshowall%3Dtrue

Rasmussen S, Allentoft ME, Nielsen K, Orlando L, Sikora M, Sjögren KG, Pedersen AG, Schubert M, Van Dam A, Kapel CM, Nielsen HB, Brunak S, Avetisyan P, Epimakhov A, Khalyapin MV, Gnuni A, Kriiska A, Lasak I, Metspalu M, Moiseyev V, Gromov A, Pokutta D, Saag L, Varul L, Yepiskoposyan L, Sicheritz-Pontén T, Foley RA, Lahr MM, Nielsen R, Kristiansen K, Willerslev E. Early divergent strains of Yersinia pestis in Eurasia 5,000 years ago. Cell. 2015 Oct 22;163(3):571-82. doi: 10.1016/j.cell.2015.10.009. Epub 2015 Oct 22. PMID: 26496604; PMCID: PMC4644222 (PubMed) https://pubmed.ncbi.nlm.nih.gov/26496604/

[34] Karmin M, Saag L, Vicente M, Wilson Sayres MA, Järve M, Talas UG, Rootsi S, Ilumäe AM, Mägi R, Mitt M, Pagani L, Puurand T, Faltyskova Z, Clemente F, Cardona A, Metspalu E, Sahakyan H, Yunusbayev B, Hudjashov G, DeGiorgio M, Loogväli EL, Eichstaedt C, Eelmets M, Chaubey G, Tambets K, Litvinov S, Mormina M, Xue Y, Ayub Q, Zoraqi G, Korneliussen TS, Akhatova F, Lachance J, Tishkoff S, Momynaliev K, Ricaut FX, Kusuma P, Razafindrazaka H, Pierron D, Cox MP, Sultana GN, Willerslev R, Muller C, Westaway M, Lambert D, Skaro V, Kovačevic L, Turdikulova S, Dalimova D, Khusainova R, Trofimova N, Akhmetova V, Khidiyatova I, Lichman DV, Isakova J, Pocheshkhova E, Sabitov Z, Barashkov NA, Nymadawa P, Mihailov E, Seng JW, Evseeva I, Migliano AB, Abdullah S, Andriadze G, Primorac D, Atramentova L, Utevska O, Yepiskoposyan L, Marjanovic D, Kushniarevich A, Behar DM, Gilissen C, Vissers L, Veltman JA, Balanovska E, Derenko M, Malyarchuk B, Metspalu A, Fedorova S, Eriksson A, Manica A, Mendez FL, Karafet TM, Veeramah KR, Bradman N, Hammer MF, Osipova LP, Balanovsky O, Khusnutdinova EK, Johnsen K, Remm M, Thomas MG, Tyler-Smith C, Underhill PA, Willerslev E, Nielsen R, Metspalu M, Villems R, Kivisild T. A recent bottleneck of Y chromosome diversity coincides with a global change in culture. Genome Res. 2015 Apr;25(4):459-66. doi: 10.1101/gr.186684.114. Epub 2015 Mar 13. PMID: 25770088; PMCID: PMC4381518, (PubMed) https://pubmed.ncbi.nlm.nih.gov/25770088/

[35] Effective population size over a generation (Ne) or over a reproductive cycle (Nb) and the adult census size (Nc) are important parameters in both conservation and evolutionary biology.

The male effective population size (Nm) refers to the number of males actively contributing to reproduction within a population. It is crucial factor in understanding genetic diversity and the impact of genetic drift. The formula used to calculate the overall effective population size (Ne) incorporates both Nm (male breeders) and Nf (female breeders): Ne = (4NmNf) / (Nm + Nf). 

Nm represents the number of males that are actually breeding and passing on their genes to the next generation. It’s not just the total number of males in the population, but rather the reproductive potential of the male.

Ferchaud, AL., Perrier, C., April, J. et al. Making sense of the relationships between Ne, Nb and Nc towards defining conservation thresholds in Atlantic salmon (Salmo salar). Heredity117, 268–278 (2016). https://doi.org/10.1038/hdy.2016.62

Waples RS. What Is Ne, Anyway? J Hered. 2022 Jul 23;113(4):371-379. doi: 10.1093/jhered/esac023. PMID: 35532202 https://pubmed.ncbi.nlm.nih.gov/35532202/

Effective Population Size, Wikipedia, This page was last edited on 10 February 2025, https://en.wikipedia.org/wiki/Effective_population_size

Kliman, R., Sheehy, B. & Schultz, J. (2008) Genetic Drift and Effective Population Size. Nature Education 1(3):3, found at Scitable: https://www.nature.com/scitable/topicpage/genetic-drift-and-effective-population-size-772523/

[36] Guyon, L., Guez, J., Toupance, B. et al. Patrilineal segmentary systems provide a peaceful explanation for the post-Neolithic Y-chromosome bottleneck. Nat Commun 15, 3243 (2024). https://doi.org/10.1038/s41467-024-47618-5

CNRS, Social change may explain decline in genetic diversity of the Y chromosome at the end of the Neolithic period, 24 Apr 2024, Phys.Org, https://phys.org/news/2024-04-social-decline-genetic-diversity-chromosome.html

Zeng, T.C., Aw, A.J. & Feldman, M.W. Cultural hitchhiking and competition between patrilineal kin groups explain the post-Neolithic Y-chromosome bottleneck. Nat Commun9, 2077 (2018). https://doi.org/10.1038/s41467-018-04375-6

[37] Guyon, L., Guez, J., Toupance, B. et al. Patrilineal segmentary systems provide a peaceful explanation for the post-Neolithic Y-chromosome bottleneck. Nat Commun 15, 3243 (2024). https://doi.org/10.1038/s41467-024-47618-5

[38] Ibid

[39] Zeng, T.C., Aw, A.J. & Feldman, M.W. Cultural hitchhiking and competition between patrilineal kin groups explain the post-Neolithic Y-chromosome bottleneck. Nat Commun9, 2077 (2018). https://doi.org/10.1038/s41467-018-04375-6

[40] Guyon, L., Guez, J., Toupance, B. et al. Patrilineal segmentary systems provide a peaceful explanation for the post-Neolithic Y-chromosome bottleneck. Nat Commun 15, 3243 (2024). https://doi.org/10.1038/s41467-024-47618-5

CRNS, Social change may explain decline in genetic diversity of the Y chromosome at the end of the Neolithic period, 24 April 2024, Phys.Org, https://phys.org/news/2024-04-social-decline-genetic-diversity-chromosome.html

[41] Banning, E. (2002). Aceramic Neolithic. In: Peregrine, P.N., Ember, M. (eds) Encyclopedia of Prehistory. Springer, Boston, MA. https://doi.org/10.1007/978-1-4615-0023-0_1

[42] Brami, Maxime, and Barbara Horejs, editors. The Central/Western Anatolian Farming Frontier: Proceedings of the Neolithic Workshop Held at 10th ICAANE in Vienna, April 2016. 1st ed., Austrian Academy of Sciences Press, 2019. JSTOR, https://doi.org/10.2307/j.ctvvh866f. Accessed 5 May 2025. https://www.jstor.org/stable/j.ctvvh866f

Kilinc, Gulsah & Koptekin, Dilek & Atakuman, Çiğdem & Sümer, Arev & Dönertaş, Handan & Yaka, Reyhan & Bilgin, Can & Büyükkarakaya, Ali & Baird, Douglas & Altınışık, Ezgi & Flegontov, Pavel & Götherström, Anders & Togan, İnci & Somel, Mehmet. (2017). Archaeogenomic analysis of the first steps of Neolithization in Anatolia and the Aegean. Proceedings of the Royal Society B: Biological Sciences. 284. 10.1098/rspb.2017.2064 https://www.researchgate.net/publication/321213055_Archaeogenomic_analysis_of_the_first_steps_of_Neolithization_in_Anatolia_and_the_Aegean

Clare, Lee & Weninger, Bernhard. (2014). The Dispersal of Neolithic Lifeways: Absolute Chronology and Rapid Climate Change in Central and West Anatolia, in The Neolithic in Turkey. Vol. 6. 10500-5200 BC: Environment, Settlement, Flora, Fauna, Dating, Symbols of Belief, with Views from North, South, East, and West (pp.1-65)
Publisher: Archaeology & Art Publications
Editors: M. Özdoğan, N. Başgelen, P. Kuniholmhttps://www.researchgate.net/publication/278156841_The_Dispersal_of_Neolithic_Lifeways_Absolute_Chronology_and_Rapid_Climate_Change_in_Central_and_West_Anatolia

Episode 17: Ceramic Neolithic Anatolia, 12 Jun 2021, Pre-History Podcast, https://prehistorypodcast.com/2021/06/12/episode-17-ceramic-neolithic-anatolia/

[43] Özdoğa, Mehmet, Archaeological Evidence on the Westward Expansion of Farming Communities from Eastern Anatolia to the Aegean and the Balkans, Current Anthropology, Volume 52, Number S4 October 2011, DOI https://doi.org/10.1086/658895

[44] Erkan Fidan, Savaş Sarıaltun, Turhan Doğan, Sezer Seçer-Fidan, Erhan İlkmen, Radiocarbon Dating Evidence and Cultural Sequencing in Chronology of Neolitchic Settlement at Bilecik-Bahçelievr from Northwest Anatlia, Mediterranean Archaeology and ArchaeometryVol. 22, No 3, (2022), pp. 133-148,DOI:10.5281/zenodo.7306042 https://www.maajournal.com/index.php/maa/article/view/811/729

Fikirtepe Culture (Pre-Pottery Neolithic A-B) (Anatolia), The History Files, https://www.historyfiles.co.uk/KingListsMiddEast/CulturesFikirtepe.htm

[45] Harun Oy, New Survey and Typological Study of Prehistoric Wares of Dutluca Region, Uşak, Turkey, Mediterranean Archaeology and ArchaeometryVol. 21, No 2, (2021), pp. 69-92, https://www.maajournal.com/index.php/maa/article/view/523/453

[46] Mehmet Özdogǎn, Archaeological Evidence on the Westward Expansion of Farming Communities from Eastern Anatolia to the Aegean and the Balkans, Current Anthropology, 52, Supplimenet 4, Oct 11, 2011, S. 415 – S430https://www.journals.uchicago.edu/doi/pdfplus/10.1086/658895

For a similar map that does not include Greece and the most of the Southern Balkans:

Illustration Twelve: Core Areas of Neolitic Formation in Anatolia Region

Click for Larger View | Source: Mehmet Özdogǎn, Archaeological Evidence on the Westward Expansion of Farming Communities from Eastern Anatolia to the Aegean and the Balkans, Current Anthropology, 52, Supplimenet 4, Oct 11, 2011, S. 415 – S430 https://www.journals.uchicago.edu/doi/pdfplus/10.1086/658895

Maxime Brami and Volker Heyd, Fig. 16a. Expansion of the DFBW horizon to the region of Marmara leading to the emergence of the ‚Archiac Fikirtepe tradition, The origins of Europe’s first farmers: The role of Hacılar and Western Anatolia, fifty years on, Oct 2011, Praehistorische Zeitschrift 86(2):Page 188, https://www.researchgate.net/publication/262605652_The_origins_of_Europe’s_first_farmers_The_role_of_Hacilar_and_Western_Anatolia_fifty_years_on

[47] Michael Rosenberg, Thomas R. Rocek, Socio-political organization in the Aceramic Neolithic of southwestern Asia: The complex evolution of socio-political complexity, Journal of Anthropological Archaeology, Volume 54, 2019, Pages 17-30, ISSN 0278-4165, https://doi.org/10.1016/j.jaa.2019.01.006.
(https://www.sciencedirect.com/science/article/pii/S0278416518301405 )

[48] Twiss KC, Bogaard A, Haddow S, Milella M, Taylor JS, Veropoulidou R, Kay K, Knüsel CJ, Tsoraki C, Vasić M, Pearson J, Busacca G, Mazzucato C, Pochron S. “But some were more equal than others:” Exploring inequality at Neolithic Çatalhöyük. PLoS One. 2024 Sep 6;19(9):e0307067. doi: 10.1371/journal.pone.0307067. PMID: 39240951; PMCID: PMC11379307, (MedPub) https://pmc.ncbi.nlm.nih.gov/articles/PMC11379307/

Twiss KC, Bogaard A, Haddow S, Milella M, Taylor JS, Veropoulidou R, et al. (2024) “But some were more equal than others:” Exploring inequality at Neolithic Çatalhöyük. PLoS ONE 19(9): e0307067. https://doi.org/10.1371/journal.pone.0307067

[49] Starčevo–Körös–Criș culture, Wikipedia, This page was last edited on 18 February 2025, https://en.wikipedia.org/wiki/Starčevo–Körös–Criș_culture

Hay, Maciamo, Starčevo–Kőrös–Criș culture (c. 6200-4500 BCE), Eupedia, https://www.eupedia.com/genetics/starcevo_culture.shtml

Prehistoric Europe, Wikipedia, This page was last edited on 15 April 2025, https://en.wikipedia.org/wiki/Prehistoric_Europe

[50] Karanovo culture, Wikipedia, This page was last edited on 23 March 2025, https://en.wikipedia.org/wiki/Karanovo_culture

Prehistoric Europe, Wikipedia, This page was last edited on 15 April 2025, https://en.wikipedia.org/wiki/Prehistoric_Europe

[51] David W. Anthony and Jennifer Y. Chi , The Lost World of Old Europe The Danube Valley, 5000–3500 bc, Princeton: Princeton University Press, 2010 https://e-edu.nbu.bg/pluginfile.php/586999/mod_resource/content/1/Anthony%20et%20al%20ed_2010_The%20Lost%20World%20of%20Old%20Europe%20Catalogue.pdf

Prehistoric Europe, Wikipedia, This page was last edited on 15 April 2025, https://en.wikipedia.org/wiki/Prehistoric_Europe

Starčevo–Körös–Criș culture, Wikipedia, This page was last edited on 18 February 2025, https://en.wikipedia.org/wiki/Starčevo–Körös–Criș_culture

Hay, Maciamo, Starčevo–Kőrös–Criș culture (c. 6200-4500 BCE), Eupedia, https://www.eupedia.com/genetics/starcevo_culture.shtml

Karanovo culture, Wikipedia, This page was last edited on 23 March 2025, https://en.wikipedia.org/wiki/Karanovo_culture

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

Karanovo culture, Wikipedia, This page was last edited on 23 March 2025, https://en.wikipedia.org/wiki/Karanovo_culture

Hamangia culture, Wikipedia, This page was last edited on 17 June 2024, https://en.wikipedia.org/wiki/Hamangia_culture

Cucuteni–Trypillia culture, Wikipedia, This page was last edited on 7 April 2025, https://en.wikipedia.org/wiki/Cucuteni–Trypillia_culture

Gumelnița culture, Wikipedia, This page was last edited on 2 April 2025, https://en.wikipedia.org/wiki/Gumelnița_culture

Gumelnița–Kodžadermen-Karanovo VI complex, Wikipedia, This page was last edited on 5 May 2024, https://en.wikipedia.org/wiki/Gumelnița–Kodžadermen-Karanovo_VI_complex

See also

History of Burgaria, Wikipedia, This page was last edited on 15 April 2025, https://en.wikipedia.org/wiki/History_of_Bulgaria

Alexandros Tsoupas, Carlos S. Reyna-Blanco, Claudio S. Quilodrán, Jens Blöcher, Maxime Brami, Daniel Wegmann, Joachim Burger, Mathias Currat, Local increases in admixture with hunter-gatherers followed the initial expansion of Neolithic farmers across continental Europe, bioRxiv, 12 Jun 2024, doi: https://doi.org/10.1101/2024.06.10.598301 , https://www.biorxiv.org/content/10.1101/2024.06.10.598301v1.full.pdf

[52] Cucuteni–Trypillia culture, Wikipedia, This page was last edited on 7 April 2025, https://en.wikipedia.org/wiki/Cucuteni–Trypillia_culture

[53] Cucuteni–Trypillia culture, Wikipedia, This page was last edited on 7 April 2025, https://en.wikipedia.org/wiki/Cucuteni–Trypillia_culture

[54] Karmin M, et al , A recent bottleneck of Y chromosome diversity coincides with a global change in culture. Genome Res. 2015 Apr;25(4):459-66. doi: 10.1101/gr.186684.114. Epub 2015 Mar 13. PMID: 25770088; PMCID: PMC4381518, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC4381518/

Immel, A., Țerna, S., Simalcsik, A. et al. Gene-flow from steppe individuals into Cucuteni-Trypillia associated populations indicates long-standing contacts and gradual admixture.Sci Rep 10, 4253 (2020). https://doi.org/10.1038/s41598-020-61190-0

[55] Guyon, L., Guez, J., Toupance, B. et al. Patrilineal segmentary systems provide a peaceful explanation for the post-Neolithic Y-chromosome bottleneck. Nat Commun 15, 3243 (2024). https://doi.org/10.1038/s41467-024-47618-5

[56] The enigma of G-PF3345 (U1, CTS342 and L497), 3 Sep 2018, G2a, YDNA Haplogroups, Population Genetics, Forums, Eupedia, https://www.eupedia.com/forum/threads/the-enigma-of-g-pf3345-u1-cts342-and-l497.37040/

[57] Ibid

[58] 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

[59] 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

[60] 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

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/

Bickle P. Thinking Gender Differently: New Approaches to Identity Difference in the Central European Neolithic. Cambridge Archaeological Journal. 2020;30(2):201-218. doi:10.1017/S0959774319000453

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

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/

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

[63] Biermann, Eric, When did Eternity End? The So Called Downfall of Linear Pottery Culture, 31 Aug 2016,22nd Annual Meeting of the EAA, Vilnuis, Lithuania, https://youtu.be/YGrZ67mx0tY

[64] 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/

Bickle P. Thinking Gender Differently: New Approaches to Identity Difference in the Central European Neolithic. Cambridge Archaeological Journal. 2020;30(2):201-218. doi:10.1017/S0959774319000453

[65] 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/