SNPs – Griffis Family

April 15, 2025April 15, 2025

Looking at the Griff(is)(es)(ith) Patrilineal Line of Descent Through the YDNA G Haplogroup Phylogenetic Tree – Part One

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 story also touches on two historical periods in the migratory path for the family patrilineal line where the lineage experienced two relatively large time gaps between known haplogroups in the ancestors’ European migratory path. Given its length, I have divided the story into two parts.

G Haplogroup European Migration in Context of European Waves of Haplogroup Migration

Illustration one provides a map depicting the general migrations of three successive waves of humans into Europe that formed the genetic ‘melting pot’ of the European population today. The peopling of Europe involved three major demographic waves that reshaped its genetic landscape: indigenous Western Hunter-Gatherers (WHG), Neolithic farmers migrating from Anatolia (Early European Farmers – EEF), and Yamnaya pastoral Western Steppe Herders (WSH) from the Pontic-Caspian region. Each group left distinct genetic markers in mitochondrial (mtDNA) and Y-chromosomal haplogroups, reflecting their origins, migrations, and interactions.

Illustration One: The Three Waves of Human Migration in Europe

The three waves of European settlement each contributed distinct haplogroup signatures. The WHG left a legacy of haplogroups mtDNA U5 and YDNA I2, while EEF introduced mtDNA N1a and YDNA G2a. The WSH migrations catalyzed a major YDNA chromosomal shift to YDNA R1b/R1a and introduced I, U2, and T1 mtDNA. The mtDNA haplogroup H’s rise to dominance (more than 40 percent in modern Europeans) reflects complex interactions with other groups and was initially rare among EEF. [1] It expanded through the admixture between Steppe groups and Neolithic survivors .

These and other genetic turnovers underscore Europe’s dynamic prehistory, where migration, cultural exchange, and demographic replacement shaped its biological and cultural landscapes. The intersection and admixture of these three waves of genetic YDNA groups influenced and partly explain the shape of the phylogenetic tree associated with the Griff(is)(es)(ith) patrilneal line.

Western Hunter-Gatherers (WHG)

The WHG populations, present in Europe since the Upper Paleolithic (~45,000–12,000 BCE), carried mtDNA haplogroups U5, U4, and U8, which dominated their genetic profile . These lineages were nearly exclusive to pre-Neolithic foragers, with U5 being the most prevalent (60–90 percent of WHG mtDNA). The absence of haplogroups like H, J, or T in early WHG underscores their genetic distinction from later arrivals . [2]

WHG males primarily belonged to haplogroup I2, particularly subclades like I2a1b, which persisted in isolated pockets into the Neolithic period . [3] Eastern Hunter-Gatherers (EHG), a related but distinct group from Russia, carried R1a and R1b lineages, foreshadowing their later prominence in Steppe populations. [4]

Early European Farmers (EEF)

Early European Farmers (EEF) from Anatolia introduced a “Neolithic package” of mtDNA haplogroups: N1a, T2, K, J, HV, V, W, and X . [5] N1a, rare in modern Europeans (less than 0.2 percent), was a signature of the Linear Pottery Culture (LBK, ~5500 BCE), reaching 15 percent frequency in early farmers . [6] Haplogroups H and U5, though initially scarce in Anatolian migrants, increased in frequency through admixture with WHG during the Middle Neolithic (approximately 4000 BCE). [7]

The Neolithic farmer migration into Europe, which began around 9,000 years ago, involved significant genetic contributions from populations originating in the Near East. The Y-chromosomal haplogroups associated with these early farmers primarily included G2a and its subclades, which are considered markers of the Neolithic expansion. The G2a haplogroup was predominant among early Neolithic farmers and is strongly associated with the spread of agriculture from the Near East into Europe. [8]

Haplogroup I2a was present in lower frequencies and is thought to represent a legacy of Mesolithic hunter-gatherer populations in Europe. While it was not dominant among Neolithic farmers, it occasionally appeared in mixed contexts, reflecting limited admixture with local hunter-gatherers. [9]

Western Steppe Herders (WSH) or pastoralists

The WSH Steppe pastoralists associated with the Yamnaya culture (~3300 BCE) introduced novel mtDNA lineages to Europe, including I, U2, and T1, which were absent in preceding populations . [10] These haplogroups, prevalent in Eastern European and Siberian foragers, comprised around 20 percent of Corded Ware Culture (CWC) mtDNA. CWC existed approximately between 3000 BCE and 2350 BE. [11] Later, the Bell Beaker Culture (BBC, about 2800 BCE) exhibited a dramatic shift to haplogroup H (about 48 percent), likely through integration of local farmer ancestry rather than direct Steppe origins . [12]

The Yamnaya expansion marked the rise of R1b-Z2103 and R1a-M417, which replaced roughly 90 percent of preceding farmer Y lineages in Central Europe by 2500 BCE . [13] Corded Ware males were initially R1b-dominant, but R1a-M417 became predominant by 2400 BCE, reflecting dynastic succession or founder effects . [14] In contrast, BBC males carried R1b-L51, a lineage that spread rapidly across Western Europe. [15]

There were some exceptions to this general wave of WSH genetic dominance in mtDNA migratory patterns. In the Eastern Baltic, WSH mtDNA (e.g., U4, U5a) persisted into the Late Neolithic, with only minimal Steppe ancestry (about 15 percent) detected in Corded Ware culture individuals. [16] This contrasts with Central Europe, where Steppe ancestry reached 75 percent. [17] The Narva culture (~4440–4240 BCE) carried haplogroup H without EEF migratory admixture, suggesting a pre-Neolithic haplogroup H presence in rare WHG subgroups. [18]

While Steppe ancestry permeated most of Europe by 2000 BCE, Iberian populations retained higher EEF ancestry (about 50 percent) and elevated the mtDNA haplogroup H frequencies, possibly due to later Bell Beaker migrations from the north . [19] The Basque people, with about 80 percent percent EEF ancestry, exemplify this genetic continuity. [20]

The decline of early European haplogroups like G2a2, which is my patrilineal genetic line of descent, and H2 represented a dramatic genetic replacement event that occurred during the Bronze Age (about 3300 BCE – 1200 BCE). Originally, G2a was the dominant haplogroup among Early European farmers, particularly in the Mediterranean region and Central Europe.

Today, G2a only makes up about 5-10 percent of Mediterranean European populations, with higher frequencies limited to isolated mountainous regions like the Apennines (15-25 percent), Sardinia (12 percent), Cantabria (10 percent), and the Austrian Alps (8 percent). Other regions with notable frequencies include northern Greece (Thessaly), Crete, Asturias (Spain), Auvergne (France), Switzerland, and Cyprus, where percentages approach or exceed 10%. These patterns reflect the survival of Neolithic populations in geographically isolated areas after successive waves of migration and invasion. [21]

While YDNA haplogrops R and I became dominant and the size of G haplogroup ancestors diminished over time, there is a plausible argument that the remaining G haplogroup males became enculturated in Steppe culture and were part of migratory patterns that coincided with the dominant R and I haplogroup males in the Bronze age and later. This absorbtion in successive emerging cultures and social structures resulted in a phylogenetic tree that has fewer branches and long durations between haplogrups.

The Interaction and Admixture of Early Hunter Gatherers and Early European Farmers

“How did the encounter between such disparate peoples unfold? … The answer is: in a kaleidoscope of different ways. There is no clear genetic evidence of interbreeding along the central European route until the LBK farmers reached the Rhine. And yet the groups mixed in other ways—potentially right from the beginning.“ [22]

Paleogenetic and anthropological studies revealed that early European farmers initially mixed minimally with local hunter-gatherers. However, during the Middle Neolithic period (6000 – 4000 BCE) [23], there was increased mixing predominantly involving hunter-gatherer males and farmer females. This pattern suggests a social structure where hunter-gatherer men integrated into farming communities, potentially reflecting power dynamics or social stratification. [24] The near absence of WHG-derived Y lineages (e.g., I2) in early Neolithic communities highlights strict patrilocal practices . [25] Patrilocal practices, in social anthropology, refer to a residence pattern where a married couple lives with or near the husband’s family, often his parents, after marriage. [26]

Previous analyses had suggested that the EEF were genetically different from other human groups from that time. A number of paleogenomic studies have claimed that the first farmers of Anatolia and Europe emerged from a population admixed between hunter-gatherers from Europe and the Near East. The mixing process started around 14,000 years ago, which was followed by a period of extreme genetic differentiation lasting several thousand years. [27]

Various studies have shown regional variations in how farming spread based on haplogroups admixture. In Britain, evidence suggests Aegean migrants virtually replaced the existing hunter-gatherer population with limited interbreeding around 4000 BCE. [28] By contrast, Baltic hunter-gatherers acquired farming knowledge through cultural exchange rather than genetic admixture, maintaining genetic continuity despite adopting new technologies. [29] In the Lower Danube Basin, which is the path the Griff(is)(es)(ith) ancestors had taken, paleogenomic evidence has revealed multi-generational mixing between Neolithic farmers and Mesolithic hunter-gatherers [30] A study by Joaquim Fort and others found that despite differences in dispersal patterns, the percentage of farmers who interbred with hunter-gatherers was remarkably consistent. [31]

The Continental Route of G Haplogroup Migration

Neolithic farmers along the Continental Route, which is the route taken by Griffis)(es)(ith) ancestors, had an effective population size approximately five times larger than contemporaneous hunter-gatherer groups. [32] This demographic advantage, combined with technological innovations like polished stone tools and pottery, enabled rapid territorial expansion. [33] These findings support a model of demic diffusion (population replacement) rather than cultural transmission alone. [34]

“The farmers reached the new continent by two routes: in boats via the Mediterranean and on foot along the Danube River from the Balkans into central Europe. Radiocarbon dating of archaeological sites revealed that by about 7,500 years ago, Danubian farmers were building villages in the Carpathian Basin—modern-day Slovakia, Hungary and Romania—and there they began creating a pottery culture. Archaeologists call it the Linear Pottery culture (LBK, by its German acronym, for Linearbandkeramik).

“Traveling rapidly westward across the fertile plains of what is now Germany, the LBK farmers reached the Rhine within just a few centuries, around 7,300 years ago. (See illustration two.) Fine-grained analysis of the evolution of pottery styles, along with radiocarbon dating, suggests that they practiced a form of leapfrog colonization. They took “stepwise movements with sometimes hundreds of kilometers covered, and then the landscape in between filled up,”…” [35]

Illustration Two: Major Rivers in Europe and the Path of the Griff(is)(es)(ith) YDNA Haplogroup Migration

Early Neolithic farmers along the Continental Route initially occupied ecological niches among the loess plains of river valleys distinct from hunter-gatherer territories (forested uplands). Isotopic analyses of LBK burials indicate diets heavily reliant on terrestrial protein (cattle and wheat), contrasting with hunter-gatherers’ marine/freshwater resources. [36] This ecological partitioning limited direct competition but also minimized genetic exchange during the LBK’s formative phase . [37]

By 5200 BCE, archaeological and genetic evidence points to increased interaction between the indigenous hunter gatherers and the farmers that migrated along the Dunube and Rhine and other tributaries:

Material cultural exchanges: Hunter-gatherer lithic styles (e.g., microliths) appear in LBK contexts. [38]
Genetic admixture: Analyses of ancient remains of Late LBK individuals reflect up to 15 percent hunter-gatherer genetic ancestry, with higher proportions in frontier zones like the North European Plain . [39]
Subsistence shifts: Some LBK communities incorporated wild game (aurochs, red deer) into their diets, reflecting adaptive hybridization. [40]

This “resurgence” of hunter-gatherer genetic influence suggests that indigenous populations were not wholly displaced but gradually integrated into expanding agricultural societies. Before the disappearance of hunter-gatherer lifeways in Europe, farmers and foragers coexisted for many generations and in certain instances for thousands of years. [41]

The Starčevo culture’s expansion into the Carpathian Basin (6000–5400 BCE) marked the first phase of the Continental Migratory Route. (See illustration two regarding the geographical cultural areas at the time.) These early farmers practiced small-scale agriculture, relying on emmer wheat and domesticated cattle, while maintaining limited contact with local hunter-gatherers . [42] Archaeo-botanical evidence from sites like Alsónyék-Bátaszék (Hungary) shows a gradual shift from wild to domesticated plant use, suggesting a transitional economy. However, genetic data from Starčevo-associated individuals reveal minimal hunter-gatherer ancestry (less then 5 percent), indicating that early interactions were largely cultural rather than biological . [43]

Illustration Two: European Cultures Between 5000 BCE and 4500 BCE

Around 5500 BCE, the Linear Pottery Culture (LBK) emerged in Transdanubia (western Hungary), catalyzing the Neolithic’s rapid dispersal into Central Europe. The interaction between Starčevo farmers and local hunter-gatherers in Transdanubia (Hungary) produced the LBKT (LBK Transdanubia), a hybrid culture blending ceramic traditions and subsistence strategies. Genetic data from Vráble-Veľké Lehemby (Slovakia) reveal that LBKT individuals retained about 95 percent of Anatolian ancestry, underscoring limited initial admixture. [44]

Characterized by distinctive incised pottery and longhouse settlements, LBK communities expanded at an average rate of 1 to 1.3 kilometers per year, reaching the Rhine Valley by 5300 BCE and the Paris Basin by 5000 BCE . [45] This “wave of advance” was facilitated by a combination of ecological and social cultural influences:

Loess soil exploitation: Fertile, wind-deposited soils supported intensive cereal cultivation. [46]
Riverine networks: The Danube, Elbe, and Rhine provided transport routes and ecological connectivity for the migratory routes of the G haplogroup. [47]
Social cohesion: LBK settlements exhibited uniform material culture, suggesting strong cultural transmission. [48]

Genetic studies of LBK populations reveal a near-complete absence of hunter-gatherer mitochondrial lineages (e.g., U5b), reinforcing the model of farmer-led migration . [49] However, sporadic admixture between the two is detectable in later phases, with hunter-gatherer ancestry rising to about 10 percent in Middle Neolithic groups . [50]

The continental route of Neolithic expansion was not a monolithic “wave of advance” but a complex process shaped by demographic, ecological, and cultural exchange. A demic diffusion dominated the early phases, with farmer migrations driven by demographic pressures and ecological preferences. Admixture increased regionally over time, reflecting an adaptive integration of hunter-gatherer populations with the early western farmers.

The Interaction and Admixture of Early European Farmers and the Steppe Pastoralists

The interaction between haplogroups G2a and R in Europe represents one of the most significant demographic transitions in human prehistory. The Neolithic G2a-bearing farmers who spread agriculture across Europe experienced a dramatic population replacement during the Bronze Age with the expansion of R1b lineages from the east. This transition was not a simple replacement, however, as evidenced by regional persistence of G2a in certain areas and the complex patterns of admixture seen in both ancient and modern populations.

Approximately 5,000 years ago (3000-2500 BCE) a remarkable migration from the Eurasian steppe dramatically transformed the genetic and linguistic landscape of prehistoric Europe. The Yamnaya culture, a group of nomadic pastoralists from what is now Russia and Ukraine, swept westward into Europe, bringing with them not only new technologies and cultural practices but also the ancestral forms of many modern European languages. [51]

Illustration three depicts the various migratory paths of the Steppe Pastoralists. The black lined route depicts the eastward migration starting east of Carpatian mountains as found in the Corded Ware culture, transforming into the successive cultures: Fatyanovo–Balanovo culture (2800 BCE), the Abashevo (2200 BCE), the Sintashta (2100-1900 BCE)-> Andronovo (1900-1700 BCE) and Indo-Aryans. The initial eastward migration around 3000 BCE, from the black circle on the map, started with the Afanasievo culture, possibly Proto-Tocharian culture. The north-westward migrations, around 2900 BCE) carrying Corded Ware culture, transforming into the Bell Beaker culture. The westward migration west of Carpatians continued into Hungary as Yamnaya, transforming into Bell Beaker culture. [52]

Illustration Three: Migration of Steppe Pastoralists

Genetic and archaeological research has provided compelling evidence for this migration and its profound impact on European prehistory. The Yamnaya were an early Bronze Age culture that emerged from the vast grasslands of the Eurasian steppe between the Southern Bug, Dniester, and Ural rivers in present-day Ukraine and Russia around 3300 BCE. They developed from an earlier population known as the Caucasus-Lower Volga (CLV) group, who lived between the Volga River and the Caucasus Mountains approximately 6,500 years ago . [53]

Archaeological and genetic evidence indicates the Yamnaya were not a genetically homogeneous population but rather a mixed group. Their genetic makeup suggests that the leading clans were of Eastern European hunter-gatherer (EHG) and Western European hunter-gatherer (WHG) paternal origin. [54] Additionally, between 35 percent and 50 percent of Yamnaya ancestry came from the South Caucasus-Zagros area, establishing a connection between “the Proto-Indo-European-speaking Yamnaya with the speakers of Anatolian languages“. [55]

The Yamnaya developed significant technological innovations that facilitated their mobile lifestyle and subsequent expansion. They were likely among the first people to herd on horseback, a revolutionary advancement that increased mobility and range. [56] They utilized wagons for transportation, allowing movement of goods and families across vast distances. [57] They brought metallurgy skills to Europe, contributing to technological advancement. [58] They were skilled at “harvesting the bioenergy of the Eurasian grasslands” through pastoral nomadism. [59]

One of the most striking aspects of the Yamnaya migration was its gender imbalance. Genetic studies suggest this was a predominantly male movement. The contrasting patterns of sex-specific migration suggest fundamentally different types of interactions between migrating and local populations during the the Neolithic expansion of early farmers and the Steppe migration. These findings help explain cultural transitions observed in the archaeological record, including changes in burial practices, technology, and social structures: [60]

The Neolithic expansion likely involved the movement of entire family units, suggesting a more gradual and integrative cultural spread.
The Steppe migration appears to have been a more male-dominated process, possibly involving male warriors or herders who then sought wives from local European farming populations.
Researchers have indicated an extremetly skewed sex ratio associated with the Steppe migration, estimating approximately 5-14 migrating males for every migrating female.
The research points to ongoing, primarily male migration from the Steppe to Central Europe over multiple generations.
The male-biased Steppe migration pattern was possibly connected to new technologies, particularly horses and chariots, and conquest strategies.

This shift represents not just a change in genetic markers but reflects profound differences in social organization, cultural practices, and technological adaptations that gave the R haplogroup populations competitive advantages.

Neolithic farming communities associated with haplogroup G, particularly G2a, established Europe’s first agricultural societies. Archaeological and genetic studies have revealed important aspects of their social organization: [61]

They were organized into patrilocal and patrilineal communities with stable family structures.
Evidence from sites like Gurgy ‘les Noisats’ in France shows these communities consisted of genetically connected pedigrees spanning multiple generations.
They practiced female exogamy (women marrying outside their group) and formed monogamous reproductive partnerships.
Their settlement patterns suggest relatively stable health conditions that supported high fertility rates.
These societies exhibited some social stratification, though perhaps less pronounced than later Indo-European groups.

The G2a farmers’ social structure prioritized stability and continuity, which aligned perfectly with their sedentary agricultural lifestyle. While functional, this stability may have made them less adaptable to competition with more militarily organized societies.

In contrast, the R haplogroup pastoralists from the Eurasian steppes developed a markedly different social configuration: [62]

Their social structure was strongly patriarchal and hierarchical with distinct social classes including priests, warriors, chiefs, commoners, and slaves.
Their socieities oncentrated wealth and power in male lineages and Increased social inequality linked to patrilineal inheritance systems.
Their social organization particularly was effective for warfare and expansion.
Young males were initiated into warrior-bands called kóryos, living by hunting, raiding, and pillaging other communities.
Their society was organized around patron-client relationships that facilitated military endeavors.
Exogamy practices allowed for the incorporation of foreign women into their societies, enabling demographic growth even in conquest scenarios.

This warrior-centric social organization gave R haplogroup populations significant advantages in conflict situations. The kóryos tradition in particular appears to have been a powerful vehicle for expansion, as these groups of young warriors could effectively raid sedentary farming communities.

Within just a few hundred years, the Yamnaya contributed to at least half of central Europeans’ genetic ancestry . [63] Their genetic impact varied geographically, with Northern Europeans showing stronger Yamnaya ancestry than Southern Europeans. It has been estimted that the Yamnaya genetic contribution in modern Eastern Europeans ranges from roughly 47 percent among Russians to 43 percent in Ukrainians, with Finland having the highest Yamnaya contribution in all of Europe at 50 percent. [64]

The genetic history revealed through these Y-chromosome lineages aligns with archaeological evidence for cultural transitions and provides insights into the formation of the modern European genetic landscape. While male lineages show evidence of substantial replacement, autosomal DNA reveals a more nuanced picture of admixture between different ancestral populations. This complex interplay between replacement and admixture has shaped the genetic diversity of Europe today, creating a mosaic of ancestry that reflects thousands of years of human migration, interaction, and cultural change.

The G Haplogroup Phylogenetic Tree in the Long Term Genealogical Time Layer

The shape and nature of the G haplogroup phylogenetic tree is for the most part long and narrow. The particular branch of the G haplogroup, which includes the migratory path of the Griff(is)(es)(ith) family, is characterized, with few exceptions, as a haplogroup with few subclades spread over a long periods of time.

Haplogroup G’s current European distribution is characterized by overall low frequency across Europe punctuated by geographically isolated regions of higher concentration. The haplogroup is historically associated with early Neolithic agricultural expansions from the Near East into Europe via the northern Mediterranean coastline (the Mediterranean route) and the inland route, traversing the Balkans and central Europe via the Danube and Rhine rivers and other tributaries.

The phylogenetic tree of haplogroup G in general reveals a complex history characterized by an early divergence into two major branches (G1 and G2), followed by subsequent diversification into geographically specific subclades. Its development spans from approximately 48,500 years ago to much more recent times, with significant expansion events occurring after the Last Glacial Maximum (26,000 – 20,000 years ago) and during the Neolithic period. ( 7000–1700 BCE ). [65]

The tree’s shape reflects multiple migration events and population bottlenecks rather than simple migration dispersal, with different subclades showing distinct geographic patterns. The higher diversity near the proposed homeland in West Asia/Middle East and the more restricted distribution of specific subclades in peripheral regions supports a model of outward migration from this central zone, with subsequent isolation and local evolution in different geographic regions. [66]

Haplogroup G (M201) emerged as one of two primary branches from the parent haplogroup GHIJK, with HIJK being its sibling clade. The overall shape of the G haplogroup tree can be characterized as having two primary branches that formed early in its history, with subsequent diversification into numerous subclades displaying strong geographical specificity.

The haplogroup G phylogenetic tree splits into two fundamental branches: the G1 (M285, M342) haplogroups which is less common globally but shows significant presence in specific regions and the G2 (P287) haplogroups which is the predominant branch containing most G lineages. [67]

The Significance of G-P15 and G-P303 Haplogroups

Haplogroup G2 further divides into: G2a (P15) which is the most widespread subclade in Europe and haplogroup G2b (M3115) which is found from the Middle East to Pakistan. Illustration four below provides a depiction of a phylogenetic tree for G-P15.

Illustration Four: Phylogenetic trees of haplogroup G2a and Haplogroup G2a-L140 (Shaded Arrows Depict Griff(is)(es)(ith) Line of Descent)

“(A) predominance or high frequency of the Y-chromosome haplogroup G- P15 in representatives of various Neolithic archaeological cultures of Europe with a pronounced decrease or complete absence in the subsequent cultures confirms the hypothesis on Neolithic expansion of Anatolian/Middle Eastern farmers toward Western Europe, which was followed by their displacement by such steppe ancestors as pro-Indo-European steppe nomads. [68]

Illustration Five: Archaeological Sites with Detected Y-Chromosome Haplogroup G-P15 Across Europe

Recent genetic studies have enhanced the understanding of G2a’s structure with several defined subclades showing geographic specificity for each of the major subclades. For example, haplogroup G2a-L140, who is a genetic ancestor of the Griff(is)(es)(ith) family line, and G2a-M406 predominate in northern and western Europe.

This genetic lineage offers insights into prehistoric human migrations, early agricultural expansions, and the complex genetic tapestry connecting populations across Europe, the Caucasus, and the Middle East. The overall coalescent age estimate for G-P303 is approximately 12,600 years ago, placing its emergence in the early post-glacial period. (The coalescent age is essentially the time to the most recent common ancestor (tMRCA) of a set of gene copies.) This timing is significant as it corresponds to a critical transition period between hunter-gatherer societies and the emergence of early agricultural communities.

Haplogroup G-P303 defines the most frequent and widespread G subclade globally. In Europe, G-P303 definable subgroups make up a majority of Haplogroup G persons west of Russia and the Black Sea, and small numbers are also found in North Africa. The percentage of haplogroup G among available samples from Wales is overwhelmingly G-P303. Such a high percentage is not found in nearby England, Scotland or Ireland. [69]

Table one below provides a comparison of specific YDNA Short Tandem Repeats (STRs), also known as microsatellites, that are associated with the G-P303 haplogrup. The table provides a comparison of these values with my test results. STRs are short, repetitive DNA sequences (typically 2-6 base pairs) that are found throughout the genome, and the number of repeats varies between individuals, making them useful for DNA identification and analysis. STR values represent the number of times a short, repeating DNA sequence occurs at a specific location (locus) on a chromosome.

“All G-P303 men carry the P303 or S135 SNP Y-DNA mutation. There are also some short tandem repeat (STR) findings among G-P303 men which help in subgrouping them. Many of the men have an unusual value of 13 for marker DYS388, and some have 9 at DYS568. STR marker oddities are often different in each G-P303 subgroup, and characteristic marker values can vary by subgroup. Often the values of STR markers DYS391, DYS392 and DYS393, however, are respectively 10, 11 and 14 or some slight variation on these for all G-P303 men.“ [70]

Table One : My Short Tandem repeats STR Values Associated with G-P303

Designated STR Y Chromosome Segment (DYS)	STR DYS Values Associated with G-P303	My STR DYS Values Associated with G-P303
YS388	13	13
DYS391	10	10
DYS392	11	11
DYS393	14	15
DYS594 (Welsh Association)	12	11

Source: My STY values are based on YDNA test results from the FamilyTreeDNA Y-700 test. [71]

The table indicates that my test results are identical to or one value off of the values associated with the G-P303 STR configuration.

A 2022 study systematically assessed the association between genetic variation in the male-specific region of the Y chromosome (MSY) and cardiovascular disease outcomes in Great Britain. While the primary finding of the study showed little evidence for an effect of any MSY haplogroup on cardiovascular risk in participants, an important secondary finding was that Y chromosome haplogroups carried by the British UK Biobank sampled individuals demonstrate a strong geographic structuring across Great Britain. [72]

This extensive analysis included up to 152,186 unrelated, genomically British individuals from UK Biobank. The Biobank has data on the whole genome sequencing for 500,000 participants, whole exome sequencing for 470,000 participants, genotyping (800,000 genome-wide variants and imputation to 90 million variants).

The prevalence of male-specific region of the Y chromosome haplogroups was plotted by place of birth in successively larger areas with at least 100 individuals, from wards and electoral divisions, to local authorities, to regions of England and the nations of Great Britain. Examples of geographic structuring of Y chromosome variation by place of birth of genetically British men from UK Biobank were depicted in maps of 90 haplogroups in the study. [73]

As reflected in illustrations five and six for haplogroups G-M201 and G-P15 respectively, there is a small prevalence of the G haplgroup in sourthern Wales where it has been hypothesized where the Griff(is)(es)(ith) family emigrated from when traveling to the American colonies.

Illustration Five: G-M201 Prevalence by Geographic Area

Illustration Six: G-P15 Prevalance by Geographic Area

The haplogroup G-L497 lineage, which is also a genetic ancestor of the Griff(is)(es)(ith) line. It belongs to the larger G2a branch and specifically the P303 subclade. G-L497 is a significant Y-chromosome haplogroup that represents one of the major European branches of haplogroup G. Most carriers, such as myself, have a distinctive value of 13 at the DYS388 marker (see table one above). [74]

G-L497’s subclades (e.g., L91, P303) correlate with specific migration waves. For instance, G2a-P303 spread into Central Europe during the Neolithic. G-L497 shows a distinct European concentration, particularly along the Rhine River and in alpine regions. The haplogroup reaches its highest frequencies in the Tyrolean Alps [75], where some valleys show concentrations above 40 percent. It is commonly found among populations in the United Kingdom and Ireland. [76] The G-L497 hsaplogroup is also found in various parts of Europe. High percentagesare found in northwestern Europe, with notable frequencies in Switzerland (74%), Spain (60%),France (58%), and Germany (57%). [77]

Illustration seven depicts the estimated early migratory path of ancestors genetically related to my G-Y132505 terminal haplogroup prior to migrating westward into Europe. The map depicts selected haplogroup subclades between G-M201, the begining of the G haplogroup (~26000 BCE) and G-L140 (~9050 BCE). It is noteworthy that for approximately 17,000 years, the YDNA G haplogroup lineage was located in the West Asia/Middle East area known as Anatolia. Anatolia, also known as Asia Minor, is the peninsula of land that today constitutes the Asian portion of Turkey.

Illustration Seven: Estimated Early Migratory Path of Griff(is)(es)(ith YDNA Lineage – 26000 BCE to 8950 BCE

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

Griff(is)(es)(ith) YDNA Phylogentic Line

Table Two provides the YDNA ancestral path of the Griff(is)(es)(ith) line from most recent haplogroup (G-BY211678) that started around 1400 CE to the distant past (Haplogroup A ). With the exception of four haplogroups, each haplogroup had two or three genetic decscendants that continued the genetic YDNA line. Below are explanations for each of the descriptive characteristics associated with each haplogroup.

Explanation of Columns in Table One

Column	Description
Haplogroup	The name of an ancestral haplogroup.
Age Estimate of MRCA Brith	The estimated birth date of the ancestor that is associated with the unique SNP mutation that is associated with the haplogroup.
Time Passed from Prior Haplogroup	Time elapsed between an haplogroup and its immediate ancestral haplogroup.
Number of Phylogenetic Subclades	Number of Phylogenetic subclades provides an indicator to esitmate the relative expansion of the subclade. A large number indicates a rapid expansion event.
Number of SNP Variants	If a haplogroup has many associated SNP variants, it suggests a greater level of genetic diversity and potential for sub-branching within that haplogroup, indicating a longer history and more complex evolutionary path. The number of sub-branches is, however, dependent upon archeaological and modern day DNA testing discoveries.

Table Two: Genetic Ancestral Path for My Terminal Haplogroup G-Y211678

Haplogroup	Age Estimate of MRCA Birth	Time Passed from Prior Haplogroup (Years)	Number of Phylogenetic Subclades	Number of SNP Variants
G-BY211678	1400 CE	300	2	3
G-Y132505	1100 CE	150	4	2
G-Z40857	950 CE	250	4	5
G-Y38335	700 CE	< 100	2	2
G-Z6748	650 CE	2,850	2	29
G-FGC7516	2200 BCE	< 100	6	1
G-FGC477	2250 BCE	300	2	1
G-Z727	2550 BCE	550	3	8
G-Z1817	3100 BCE	850	2	10
G-Z6901	3950 BCE	650	1	5
G-Z1900	4600 BCE	300	2	4
G-CTS9737	4900 BCE	600	1	9
G-L497	5500 BCE	3,450	2	48
G-PF3345	8950 BCE	< 100	11	1
G-Z3065	8950 BCE	< 100	2	2
G-PF3346	9000 BCE	< 100	3	1
G-L140	9050 BCE	800	3	14
G-P303	9850 BCE	2,150	3	38
G-L141	12,000 BCE	1,000	2	14
G-L30	13,000 BCE	3,000	2	47
G-L1259	16,000 BCE	< 1,000	2	7
G-P15	16,000 BCE	4,000	2	56
G-L156	20,000 BCE	4,000	2	62
G-L89 (G2)	24,000 BCE	2,000	2	32
G-M201 (G)	26,000 BCE	20,000	2	324
GHIJK-F1329	46,000 BCE	1,000	2
F-M89	47,000 BCE	16,000	2
CF-P143	63,000 BCE	1,000	2
CT-M168	64,000 BCE	22,000	2
BT-M42	86,000 BCE	35,000	2
A-V221 (A1b)	121,000 BCE	5,000	2
A-V168 (A1)	126,000 BCE	6,000	2
A-L1090 (A0T)	152,000 BCE	80,000	2
A-PR2921 (Y-Adam)	232,000 BCE	136,000	2
A000-T(Neanderthal Divergence)	368,000 BCE	337,000	2
A-0000 (Denisovan divergence)	705,000 BCE

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

If attention is focused on haplogroups that are after G-L140 (~9050 BCE), there are two notable gaps between haplogroups or most common recent ancestors in the migratory chain.

The 3,500 year Gap between G-PF3345 and G-L497: The common ancestor born around 8550 BCE, associated with the haplogroup G-PF3345, had eleven surviving descendants. The next common ancestor was associated with the G-L497 haplogroup about 3,500 years later. The time that passed from the prior ancestor to G-PF3345 was less than 100 years.
The 2,850 year Gap between G-FGC7516 and G-Z6748: The other common ancestor born around 2200 BCE, associated with haplogroup G-FGC477, had 6 surviving descendants and the prior ancestor was less than 100 years. The next genetic ancestor was associated with the genetic SNP mutation defining the G-Z6748 haplogroup 2,850 years later. Similar to haplogroup G-PF3345, the preceding genetic ancestor to G-FGC7516 was less than 100 years .

Illustration Eight depicts these two gaps in a map of the estimated migration path of the Griff(is)(es)(ith) patrilineal line. The map depicts various haplogroups along an estimated migratory path for the patrilineal family line.

A dotted arrow connects haplogroup G-PF3345 with haplogroup G-L497. As reflected in the illustration, there a large time gap between the two common descendants. It is also evident that there are no other known, documented G haplogroup descendants that migrated up the western coast of the Black sea and moved westward following the Danube River to an approximate area presently known as Tyrol, Austria. It is in this general area that the ancestor associated with the G-L497 haplogroup was born. [78]

Illustration Eight: Highlighted Haplogroup Gaps in Griff(is)(es)(ith) Migratoy Path

Between roughly 8950 BCE and 5500 BCE, the Griff(is)(es)(ith) family paternal ancestral path was a mystery. Based on geographical and environmental data, it would appear that the migratory path followed the western outline of the Back Sea and continued westward along the second longest river in Europe, what is now known as the Danube River. During this time period the haplogroup path and phylogenetic tree of descendants appears to have been long and narrow.

The other major gap between haplogroups G-FGC7516 and GZ6748 is depicted in illustration nine by an enlarged area of the map in illustration eight. The second phylogenetic gap corresponds with a northward migratory path that is found near modern day Luxemburg and Western Germany.

Illustration Nine: Estimated Migratory Path Between G-FGC7516 and G-Z6748

The estimated northward migratory path is in an area that has many rivers that may have influenced and facilitated migratory travel, as depicted in illustration ten. There are a number of major tributaries around the estimated migratory path of the Griff(is)(es)(ith paternal YDNA line: the Rhine, the Meuse, the Maas, the Waal, the Mosel, and the Neckar rivers.

Illustration Ten: Rivers in Northwestern Europe

The cultural landscape of northwestern Europe underwent dramatic transformations between 2200 BCE and 650 CE, spanning the transition from the Late Neolithic through the Bronze Age, Iron Age, Roman period, and into the early Medieval era. This period witnessed significant shifts in technology, social organization, settlement patterns, and ethnic composition, driven by a complex interplay of migration, trade, technological innovation, and political developments.

Part Two of this Story

There are a number of possible explanations for these phylogenetic gaps that are uniquely tied to demographic, social and historic influences in each of these time periods. There also are obvious methodogical explanations for each of the two historical historical gaps in phylogenetic branching. The second part of this story discusses possible explanations and the social and cultural influences in these time periods associated with the two phylogenetic gaps.

Source:

Feature Banner: The banner at the top of the story is a modification of a shapshot taken from the FamilyTreeDNA Globetrekker^TM video of the migratory path of my YDNA descendants over time. The map shows the maigratory path of selected most common recent ancestors and their respective eestimated dates of birth. In addition the map highlights two time periods where there was a signficant time period inbetween haplogroups that are discussed in the present story.

[1] Brandt G, Haak W, Adler CJ, Roth C, Szécsényi-Nagy A, Karimnia S, Möller-Rieker S, Meller H, Ganslmeier R, Friederich S, Dresely V, Nicklisch N, Pickrell JK, Sirocko F, Reich D, Cooper A, Alt KW; Genographic Consortium. Ancient DNA reveals key stages in the formation of central European mitochondrial genetic diversity. Science. 2013 Oct 11;342(6155):257-61. doi: 10.1126/science.1241844. PMID: 24115443; PMCID: PMC4039305, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC4039305/

Brotherton, P., Haak, W., Templeton, J. et al. Neolithic mitochondrial haplogroup H genomes and the genetic origins of Europeans. Nat Commun 4, 1764 (2013). https://doi.org/10.1038/ncomms2656

Early European Farmers, Wikipedia, This page was last edited on 9 March 2025, https://en.wikipedia.org/wiki/Early_European_Farmers

[2] Brandt G, Haak W, Adler CJ, Roth C, Szécsényi-Nagy A, Karimnia S, Möller-Rieker S, Meller H, Ganslmeier R, Friederich S, Dresely V, Nicklisch N, Pickrell JK, Sirocko F, Reich D, Cooper A, Alt KW; Genographic Consortium. Ancient DNA reveals key stages in the formation of central European mitochondrial genetic diversity. Science. 2013 Oct 11;342(6155):257-61. doi: 10.1126/science.1241844. PMID: 24115443; PMCID: PMC4039305, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC4039305/

Hervella M, Izagirre N, Alonso S, Fregel R, Alonso A, Cabrera VM, de la Rúa C. Ancient DNA from hunter-gatherer and farmer groups from Northern Spain supports a random dispersion model for the Neolithic expansion into Europe. PLoS One. 2012;7(4):e34417. doi: 10.1371/journal.pone.0034417. Epub 2012 Apr 25. Erratum in: PLoS One. 2012;7(7). doi:10.1371/annotation/3dac0b4f-f76e-4bc1-8559-acb41b87b02c. PMID: 22563371; PMCID: PMC3340892, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC3340892/

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

L.G. Simões, R. Peyroteo-Stjerna, G. Marchand, C. Bernhardsson, A. Vialet, D. Chetty, E. Alaçamlı, H. Edlund, D. Bouquin, C. Dina, N. Garmond, T. Günther, & M. Jakobsson, Genomic ancestry and social dynamics of the last hunter-gatherers of Atlantic France, Proc. Natl. Acad. Sci. U.S.A. 121 (10) e2310545121, https://doi.org/10.1073/pnas.2310545121 (2024).

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

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

[4] Western Steppe Herders, Wikipedia, This page was last edited on 27 March 2025, https://en.wikipedia.org/wiki/Western_Steppe_Herders

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

[5] Brandt G, et al , Ancient DNA reveals key stages in the formation of central European mitochondrial genetic diversity. Science. 2013 Oct 11;342(6155):257-61. doi: 10.1126/science.1241844. PMID: 24115443; PMCID: PMC4039305, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC4039305/

Z. Hofmanová, S. Kreutzer, G. Hellenthal, C. Sell, et al , Early farmers from across Europe directly descended from Neolithic Aegeans, Proc. Natl. Acad. Sci. U.S.A.113 (25) 6886-6891, https://doi.org/10.1073/pnas.1523951113 (2016).

Fernández E, Pérez-Pérez A, Gamba C, Prats E, Cuesta P, Anfruns J, Molist M, Arroyo-Pardo E, Turbón D. Ancient DNA analysis of 8000 B.C. near eastern farmers supports an early neolithic pioneer maritime colonization of Mainland Europe through Cyprus and the Aegean Islands. PLoS Genet. 2014 Jun 5;10(6):e1004401. doi: 10.1371/journal.pgen.1004401. PMID: 24901650; PMCID: PMC4046922, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC4046922/

[6] Brandt G, et al ,Ancient DNA reveals key stages in the formation of central European mitochondrial genetic diversity. Science. 2013 Oct 11;342(6155):257-61. doi: 10.1126/science.1241844. PMID: 24115443; PMCID: PMC4039305, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC4039305/

Brotherton, P., Haak, W., Templeton, J. et al. Neolithic mitochondrial haplogroup H genomes and the genetic origins of Europeans. Nat Commun 4, 1764 (2013). https://doi.org/10.1038/ncomms2656

[7] Brotherton, P., Haak, W., Templeton, J. et al. Neolithic mitochondrial haplogroup H genomes and the genetic origins of Europeans. Nat Commun 4, 1764 (2013). https://doi.org/10.1038/ncomms2656

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

Early European Farmers, Wikipedia, This page was last edited on 9 March 2025, https://en.wikipedia.org/wiki/Early_European_Farmers

[8] 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, https://pmc.ncbi.nlm.nih.gov/articles/PMC4389623/

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

Ainash Childebayeva, Adam Benjamin Rohrlach, Rodrigo Barquera, Maïté Rivollat, Franziska Aron, András Szolek, Oliver Kohlbacher, Nicole Nicklisch, Kurt W. Alt, Detlef Gronenborn, Harald Meller, Susanne Friederich, Kay Prüfer, Marie-France Deguilloux, Johannes Krause, Wolfgang Haak, Population Genetics and Signatures of Selection in Early Neolithic European Farmers, Molecular Biology and Evolution, Volume 39, Issue 6, June 2022, msac108, https://doi.org/10.1093/molbev/msac108

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

Ainash Childebayeva, et al., Population Genetics and Signatures of Selection in Early Neolithic European Farmers, Molecular Biology and Evolution, Volume 39, Issue 6, June 2022, msac108, https://doi.org/10.1093/molbev/msac108

[10] Brandt G, et al ,Ancient DNA reveals key stages in the formation of central European mitochondrial genetic diversity. Science. 2013 Oct 11;342(6155):257-61. doi: 10.1126/science.1241844. PMID: 24115443; PMCID: PMC4039305, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC4039305/

Juras, A., Chyleński, M., Ehler, E. et al. Mitochondrial genomes reveal an east to west cline of steppe ancestry in Corded Ware populations. Sci Rep 8, 11603 (2018). https://doi.org/10.1038/s41598-018-29914-5

Maja Krzewińska et al. ,Ancient genomes suggest the eastern Pontic-Caspian steppe as the source of western Iron Age nomads.Sci. Adv.4,eaat4457(2018).DOI:10.1126/sciadv.aat4457 , https://www.science.org/doi/10.1126/sciadv.aat4457

[11] Juras, A., Chyleński, M., Ehler, E. et al. Mitochondrial genomes reveal an east to west cline of steppe ancestry in Corded Ware populations. Sci Rep 8, 11603 (2018). https://doi.org/10.1038/s41598-018-29914-5

Wolfgang Haak, Iosif Lazaridis, Nick Patterson, Nadin Rohland, Swapan Mallic, Bastien Llamas1, Guido Brandt, et al, Massive migration from the steppe was a source for Indo-European languages in Europe, Research Letter, 2015, Nature, https://reich.hms.harvard.edu/sites/reich.hms.harvard.edu/files/inline-files/nature14317.pdf

[12] Brandt G, et al ,Ancient DNA reveals key stages in the formation of central European mitochondrial genetic diversity. Science. 2013 Oct 11;342(6155):257-61. doi: 10.1126/science.1241844. PMID: 24115443; PMCID: PMC4039305, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC4039305/

Brotherton, P., Haak, W., Templeton, J. et al. Neolithic mitochondrial haplogroup H genomes and the genetic origins of Europeans. Nat Commun 4, 1764 (2013). https://doi.org/10.1038/ncomms2656

Early European Farmers, Wikipedia, This page was last edited on 9 March 2025, https://en.wikipedia.org/wiki/Early_European_Farmers

[13] Western Steppe Herders, Wikipedia, This page was last edited on 27 March 2025, https://en.wikipedia.org/wiki/Western_Steppe_Herders

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

[14] Wolfgang Haak, Iosif Lazaridis, Nick Patterson, Nadin Rohland, Swapan Mallic, Bastien Llamas1, Guido Brandt, et al, Massive migration from the steppe was a source for Indo-European languages in Europe, Research Letter, 2015, Nature, https://reich.hms.harvard.edu/sites/reich.hms.harvard.edu/files/inline-files/nature14317.pdf

Haak W, Lazaridis I, Patterson N, Rohland N, et al , Massive migration from the steppe was a source for Indo-European languages in Europe. Nature. 2015 Jun 11;522(7555):207-11. doi: 10.1038/nature14317. Epub 2015 Mar 2. PMID: 25731166; PMCID: PMC5048219, (Pubmed) https://pmc.ncbi.nlm.nih.gov/articles/PMC5048219/

[15] Wolfgang Haak, Iosif Lazaridis, Nick Patterson, Nadin Rohland, Swapan Mallic, Bastien Llamas1, Guido Brandt, et al, Massive migration from the steppe was a source for Indo-European languages in Europe, Research Letter, 2015, Nature, https://reich.hms.harvard.edu/sites/reich.hms.harvard.edu/files/inline-files/nature14317.pdf

[16] Mittnik, A., Wang, CC., Pfrengle, S. et al. The genetic prehistory of the Baltic Sea region.Nat Commun 9, 442 (2018). https://doi.org/10.1038/s41467-018-02825-9

[17] Brotherton, P., Haak, W., Templeton, J. et al. Neolithic mitochondrial haplogroup H genomes and the genetic origins of Europeans. Nat Commun 4, 1764 (2013). https://doi.org/10.1038/ncomms2656

Daniel M. Fernandes, Alissa Mittnik, et al, The spread of steppe and Iranian-related ancestry in the islands of the western Mediterranean, Nature Ecology & Evolution, https://reich.hms.harvard.edu/sites/reich.hms.harvard.edu/files/inline-files/2020_Fernandes_NatEcolEvol_WestMediterranean_0.pdf

[18] Early European Farmers, Wikipedia, This page was last edited on 9 March 2025, https://en.wikipedia.org/wiki/Early_European_Farmers

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

[19] Brotherton, P., Haak, W., Templeton, J. et al. Neolithic mitochondrial haplogroup H genomes and the genetic origins of Europeans. Nat Commun 4, 1764 (2013). https://doi.org/10.1038/ncomms2656

Daniel M. Fernandes, Alissa Mittnik, Iñigo Olalde, et al., The spread of steppe and Iranian-related ancestry in the islands of the western Mediterranean, Nature Ecology & Evolution , https://doi.org/10.1038/s41559-020-1102-0 , https://reich.hms.harvard.edu/sites/reich.hms.harvard.edu/files/inline-files/2020_Fernandes_NatEcolEvol_WestMediterranean_0.pdf

[20] Early European Farmers, Wikipedia, This page was last edited on 9 March 2025, https://en.wikipedia.org/wiki/Early_European_Farmers

[21] Sjur Cappelen Papazian, The spread of haplogroup G2a in Europe, 2 Nov 2013, Cradle of Civilization, https://aratta.wordpress.com/2013/11/02/bell-beakers/

Hay, Maciamo, Distribution of G2a in Italy (Boattini et al.), 7 jun 2013, Forum, Eupedia, https://www.eupedia.com/forum/threads/distribution-of-g2a-in-italy-boattini-et-al.28663/

Hay, Maciamo, Origins and history of Haplogroup G2a (Y-DNA), Oct 2016, https://www.academia.edu/6173684/Origins_and_history_of_Haplogroup_G2a_Y_DNA_

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

[22] Laura Spinney’s article “When the First Farmers Arrived in Europe, Inequality Evolved” highlights several key points regarding the societal changes and the rise of inequality associated with the arrival of agriculture in Europe.

Long-term Impacts on European Society: The establishment of farming led to surplus food production, which facilitated trade and storage but also created conditions for social stratification and inequality. Communities became more complex socially and economically, laying foundations for later societal structures characterized by hierarchy and unequal resource distribution

Introduction of Agriculture and Inequality: The arrival of the first farmers in Europe, migrating from Anatolia around 7,000 BC, marked a significant shift from hunter-gatherer societies to settled agricultural communities. This transition led to increased population densities and sedentary lifestyles .

Emergence of Social Hierarchies: With agriculture came surplus food production, which allowed for accumulation of resources. This surplus facilitated the development of social hierarchies and economic disparities, as some individuals or groups could accumulate more resources than others, leading to the evolution of inequality.

Genetic Evidence of Social Stratification: Archaeogenetic studies revealed that early European farmers initially mixed minimally with local hunter-gatherers; however, during the Middle Neolithic period, there was increased mixing predominantly involving hunter-gatherer males and farmer females. This pattern suggests a social structure where hunter-gatherer men integrated into farming communities, potentially reflecting power dynamics or social stratification.

Violence and Competition: The archaeological record includes evidence of violence among early farming groups, such as the systematic massacre at Els Trocs in Spain, indicating competition for resources or territory among early agricultural communities.

Spinney, Laura, When the First Farmers Arrived in Europe, Inequality Evolved, 1 Jul 2020, Scientific American, https://www.scientificamerican.com/article/when-the-first-farmers-arrived-in-europe-inequality-evolved/

The Griff(is)(es)(ith) Patrilineal Line of Descent: The Shape and Movement of the G Phylogenetic Tree through Time

This story focuses on looking at the 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.

Y-DNA phylogenetic trees provide an effective, graphic portrayal of human genetic history and genealogy. They offer insights into paternal lineage, population migrations and a complimentary image to discuss anthropological research and genealogical connections. Phylogenetic trees are also known as an evolutionary tree, cladogram, or tree of life. [1]

The use of phylogenetic trees provide a skeletal outline of the specific evolutionary path of the patrilineal genetic line of the Griff(is)(es)(ith) family. The family genetic patrilineal line is part of Haplogroup G. The G haplogrup is a Y-chromosomal lineage originating in the eastern Anatolian-Armenian-western Iranian region. From aproximately 10,000 BCE to 3,000 BCE it was a predominant YDNA haplogrup in Europe. Thereafter, it lost its predomance and became a minorty among YDNA haplogroups in Europe.

Looking Backward in Time: The Present European Y-DNA Phylogenetic Tree

In 2013 FamilyTreeDNA (FTDNA) released the advanced Big Y test and since then the company analyzed 32,000 Y chromosomes in ultra-high resolution. This has allowed the ability to identify hundreds of thousands of unique Y chromosome mutations. In 2019, the company created the Y-700 YDNA test and detected over 500,000 unique mutations in 32,000 Big Y testers. In May 2019, the Y-DNA haplotree passed 20,000 branches. The branches are defined by over 150,000 unique mutations. [2]

Illustration one below represents a circular phylogenetic Y-DNA haplogroup tree based on the testing results of FTDNA in 2019. It is a visual representation that shows evolutionary relationships between paternal lineages. The tree structure displays branches that represent genetic mutations and divergence over time. Time flows from the center outward, with older lineages near the center and younger ones at the periphery. Each branching point represents a most recent common ancestor – a genetic mutation (SNP) that created a new haplogroup. [3] Related haplogroups are grouped together in adjacent branches, showing their evolutionary relationship. Branch length indicates genetic distance or time between the genetic mutations.

Illustration One: FamilyTreeDNA Circular Phyogenetic YDNA Tree of Haplogroups Based on the Y-700 Test Results

A review of the ‘pie chart’ or circular phylogenetic tree in illustration one reveals the predominance of the R haplogroup. Haplogroup R represents about half the men who have completed FamilyTreeDNA Y-700 DNA tests and has several major subbranches. Haplogroups I and J present roughly one third of the men tested. The Griff(is)(es)(ith) family lineage is part of haplogroup G, which is an older haplogroup with fewer branches and fewer test results.

While this circular phylogenetic tree represents the 2019 population of FTDNA Y-700 DNA tests kits, it has a vague proportional resemblance of the YDNA composition of Europe.

“We know that two-thirds of all European men descend from just three ancestors who lived in the late Neolithic.” [4]

This quote is an attention grabber. It has been quoted in a number of genealogical sources. In most regions of Europe, the Neolithic period generally ended around 3000 BCE, marking the transition to the Bronze Age. The exact time frame can vary depending on the geographical location with some areas seeing the Neolithic last until around 2000 BC. [5]

This remarkable genetic pattern emerged during a massive population explosion that occurred across Europe during the Bronze Age, spanning from the Balkans to the British Isles. The population expansion occurred between 2,000 and 4,000 years ago, particularly affecting males across a continuous region from Greece to Scandinavia. These dominant males, likely associated with Bronze Age cultures, established lineages that became prevalent throughout European populations. [6]

This YDNA genetic legacy differs from patterns seen in mitochondrial DNA which is passed down through mothers. Research of mtDNA genetic patterns shows much older population growth patterns, suggesting this was a male-specific phenomenon tied to Bronze Age social structures. [7]

Going back to the original quote regarding those three ancestors, the statement requires some clarification. Genetic studies show that approximately sixty-four percent of European men can trace their Y-chromosome lineages back to just three male ancestors who lived between 3,500 and 7,300 years ago. These haplogroup lineages are identified as I1, R1a, and R1b and are identified in illustration five by three ‘standing male’ symbols. [8]

By counting the number of mutations that have accumulated within each branch over the generations, it is estimated that these three men lived at different times between 3,500 and 7,300 years ago. The lineages of each seem to have exploded in the centuries following their lifetimes to dominate Europe. The Bronze age is identified by a dotted elliptical circle in the illustration. Within that enclircled time era, the idenification of a proliferaion of lineages is evident in the I and R haplogroups.

Illustration Two: Phylogeny and Geographical Distribution of European Lineages

The spread of these Y-chromosome patterns depicted in illustration two may be linked to the influence of the Yamnaya people. They were nomadic, pastoral herders from the steppes of modern-day Ukraine and Russia. They entered Europe around 4,500 years ago. They brought with them technological innovations including horses, the use of wheel driven transportation, and distinctive burial practices. Dominant males linked with these cultures could be responsible for the Y chromosome patterns we see today. [9]

The complete genetic heritage of modern Europeans is complex, involving at least three distinct ancestral populations and represented by a number of YDNA and mtDNA Haplogroups: West European Hunter-Gatherers, Ancient North Eurasian and Early European Farmers. This genetic mixing occurred within the last 7,000 years, creating the modern European gene pool. [10]

Overview of the Migratory Path

As a background to discussing the patrilineal line of descent, the video below is an animated version of the estimated migratory path of the genetic Y-DNA descendants of the Griff(is)(es)(ith) family line. It is a singular path based on my Y-700 YDNA test results. It starts with the root Y-DNA source in Africa, often referred to as “Y-chromosomal Adam,” the most recent common ancestor of all living males. [11]

Animated Video of Estimated Migratory YDNA Path for the Griff(is)(es)(ith) Paternal Line

Source: Migratory Rendition for Griffis Family Y-DNA Migratory path, Globetrekker, FamilyTreeDNA

Click to view larger version of the video.

The animated video provides an inuitive rendition of over 200,000 years of the successive mutations in Y-DNA for the family paternal line. It provides a graphic portrayal of the general path of migration that ultimately led to the English Isle. The animation depicts lands that are now submerged (e.g. Doggerland [12] ) and the extent of the ice age in context of the migration. Illustration one below is a graphic portrayal the migratory path of the G haplogroup starting around 26,000 BCE.

Illustration One: Snapshot of Migratory Path of G Haplogroup and Griff(is)(es)(ith) Family Descendants

Haplogroup G-M201 likely originated in a region spanning eastern Anatolia, Armenia, and western Iran around 26,000 BCE.. The earliest G-M201 carriers were linked to pre-Neolithic populations, but its diversification in other subclades accelerated during the Neolithic transition (around 10,000 BCE). The G-P303 sub-clade, which accounts for the majority of European G lineages, diverged during this period, with sub-clades like G-L497 (Europe-specific) and G-U1 (Near Eastern/Caucasus) reflecting later regional adaptations. [13]. Haplogroups G2a and J-M172, which originated in Anatolia, spread westward alongside early farming communities. [14]

Haplogroup G2a spread across Europe primarily through the Neolithic agricultural expansion from the Near East (Anatolia) into Europe, roughly between 9,000 and 5,000 years ago. This migration involved early farming communities moving westward, introducing agriculture, domesticated animals, and pottery cultures into regions previously inhabited by hunter-gatherers. 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. [15]

The Two Routes of G Haplogroup Migration

As depicted in illustation two below, the spread of the G haplogroup occurred via two main routes: the Mediterranean Coastal Route (“Maritime Route”) and the Central European Inland Route (“Danubian Route”).

Illustration Two: The Two Main Routes of Migration for Neolithic Farmers

The Mediterranean route took early Neolithic farmers carrying haplogroup G2a along the Mediterranean coastline, establishing settlements in Greece, Italy, southern France, Spain, and Portugal. This migration is associated with the Cardium Pottery culture, characterized by pottery decorated with shell impressions. Ancient DNA evidence from Neolithic sites in southern France (such as the Treilles group around 3000 BCE) confirms a high prevalence of G2a ( individuals who descended from populations originating in Anatolia or the Aegean region. [16]

Another major route was inland via the Danube River valley into Central Europe. This is the route that the Griff(is)(es)(ith Paternal genetic line of descent took in migranting westward in Europe. This dispersal is associated with the Linear Pottery culture (LBK) (approximately 5500–4500 BCE), which introduced agriculture to Central Europe. Ancient DNA analyses of LBK archaeological sites in Germany and Hungary show a high frequency of haplogroup G2a among early farmers. [17]

By 7000 BCE, these practices spread northwestward into southeastern Europe, marking the start of the Continental Route . The Starčevo culture (6000–5400 BCE) in present-day Serbia and Hungary served as the initial bridge between Anatolian farmers and the Danube Basin, establishing agro-pastoral communities that later influenced the LBK . [18]

The G haplogroup associated with this ‘Danbian route’ in central Europe shows a frequency peak in the Danube basin associated with the G-L497 haplogroup, aligning with the Linear Pottery Culture (LBK) expansion. [19] The European origin of G-L497 makes it particularly valuable for tracing secondary migration patterns, such as the Griff(is)(es)(ith) paternal line, and population movements within Europe following the initial Neolithic expansion.

G Haplogroup Decline, Absorption and Refuge

Despite its widespread initial distribution, along with the J Haplogroup, during Europe’s Neolithic period, haplogroup G2a significantly declined in frequency after 3000 BCE due to migrations of pastoralist populations from the Eurasian steppe (such as the Yamnaya culture), who carried different Y-DNA haplogroups like R1b and R1a. These migrations largely replaced or assimilated earlier farming populations. [20]

The R haplogroup pastoralists expanded through the Pontic-Caspian steppe corridor, moving westward into Europe from their eastern origins. These steppe populations were genetically distinct from both European hunter-gatherers and early farmers. The expansion of these pastoralist groups led to massive population turnover in Europe, with substantial genetic input from steppe populations arriving after 3000 BCE. [21]

While many G2a lineages were largely replaced by Indo-European expansions, some G2a-L140 subclades appear to have been assimilated into Proto-Indo-European societies associated with the R haplogroups. These lineages, including certain L497-derived groups, joined R1b and R1a tribes in their subsequent migrations. This suggests a complex interaction between the descendants of Neolithic farmers and the expanding Indo-European populations rather than simple replacement. [22]

These “Indo-Europeanized G2a lineages”, such as the Griff(is)(es)(ith) line, belonged to deep clades of G2a-L140, including subclades like L13 and Z1816. While the original Neolithic G2a populations were dramatically reduced, some were incorporated into the expanding Indo-European groups, allowing certain G-L497 lineages to spread alongside R1a and R1b haplogroups during later migrations.

Today, haplogroup G2a descendants remain present at lower frequencies throughout Europe but have higher concentrations in isolated regions like Sardinia and parts of the Caucasus, reflecting remnants of these ancient Neolithic expansions. The Griffis)(es)(ith) paternal line is part of this minorty haplogroup in modern times.

Haplogroups and Phylogenetic Trees

Y-DNA haplogroups serve as markers of historical population movements. A haplogroup is a group of people who share a common ancestor and similar genetic markers. The Y chromosome’s lack of recombination allows SNPs (single nucleotide polymorphisms) to accumulate linearly over generations, making them valuable markers for tracing paternal lineages.

Human phylogenetics is the study of evolutionary relationships between ancient and present humans based on their genetic material, specifically through DNA and RNA sequencing. Phylogenetic relationships are typically visualized through phylogenetic trees, which use branches and nodes to show the chronology of genetic mutations. These trees can be either rooted, showing a hypothetical common ancestor, or unrooted, making no assumptions about ancestral lines. [23]

“Classifying the accumulated SNPs generation by generation make it possible to retrace the genealogical tree of humanity with great accuracy, to detect patterns in the distribution of shared historical lineages and to retrace historical migrations of male lineages.” [24]

Y-DNA phylogenetic trees are visual representations of the evolutionary relationships between different paternal lineages in human populations based on mutations in the Y chromosome. These trees illustrate the hierarchical structure of Y-DNA haplogroups, which are groups of men sharing specific mutations on their Y chromosome inherited from common paternal ancestors.

“Paleolithic lineages that underwent serious population bottlenecks for thousands of years sometimes have a series of over one hundred defining SNPs or SNP variants (e.g. haplogroups G and I1 each have over 300 defining SNPs). Generally speaking the number of accumulated SNPs between a haplogroup and its direct subclade correlates roughly to the number of generations elapsed.” [25]

The average number of years between Y-chromosomal SNP mutations is a parameter for estimating timelines in genetic genealogy, population genetics, and anthropological studies. Based on current research and commercial testing methodologies, this interval typically ranges from 83 to 144 years per SNP, depending on the sequencing technology, genomic regions analyzed, and mutation rate calculations. [26]

Branch lengths in a YDNA phylogenetic tree can be interpreted as measures of time, but there is significant scientific debate about the exact temporal relationships. [27] In phylogenetic studies (the study of evolutionary relationships between human remains or tests based on genetic material), branch lengths are considered proportional to time when evolution rates are uniform across lineages. [28] For Y-chromosomes, this has allowed researchers to create phylogenies where branch lengths can be used to estimate the timing of population divergences. [29]

Y-DNA Phylogenetic Trees

The phylogenetic tree starts with a root, often referred to as “Y-chromosomal Adam” [30], the most recent common ancestor of all living males. Haplogroups are labeled with letters A through T, with further subclades denoted by numbers and lowercase letters. The Y Chromosome Consortium (YCC) developed a naming system for major haplogroups and their subclades. [31]

Illustration Three: Major Clades of Y-DNA Phylogenetic Tree

Phylogenetic trees contextualize these haplogroups within historical and geographical frameworks, revealing how subclades diverged during key migratory periods. The combination of Y-DNA trees with archaeological findings has clarified debates over human migratory patterns. Each branch represents a distinct lineage defined by specific single-nucleotide polymorphisms (SNPs). The tree’s depth indicates the time since divergence with deeper branches representing older lineages. New mutations are continually discovered, leading to regular updates and the increased resolution of the tree. [32]

Illustration Four: Chronological Development of Main Western Eurasian Y-DNA Haplogroup Subclades from the Late Paleolithic to the Iron Age

Y-DNA phylogenetic trees provide a number of advantages for genealogical studies, forensic applications and population genetics. They can resolve paternal lineages and surname correlations, validate and extend surname clusters, enhance foresensc and kinship analysis, advance methodological innovations, and reconstruct ancient migrations and population histories. [33]

These trees can integrate short tandem repeats (STRs) and SNPs to resolve relationships across both recent, mid range and deep historical time scales. By dating branch points using mutation rates, researchers estimate the timing of population splits. Classiying SNPs and STRs into a genealogical order is known as phylogenentics. [34]

Y-DNA phylogenetic trees excel in connecting individuals who share recent common ancestors through STR markers, which mutate relatively quickly, and deeper ancestral links through slower-mutating SNPs. For example, STR-based clusters (e.g., 37-marker or 111-marker STR haplotypes) can identify related individuals within a genealogical timeframe in the last 500 years, while SNP-defined haplogroups (for example, the G-L497 haplogroup) trace lineage splits dating to the Neolithic or Bronze Age. This dual resolution allows surname projects to corroborate paper trails with genetic evidence, particularly for patrilineal lines where records are sparse in the short term and mid range genealogical time layers. [35]

The Most Recent Common Ancestor and Phylogenetic Trees

The ‘nodes’ in phylogenetic trees represent estimated birth dates of the most recent common ancestors for subsequent lineages. The ages of the most recent common ancestors (tMRCA) in Y-DNA phylogenetic trees are calculated primarily through statistical methods that incorporate genetic data and historical information.

Rather than focus on the order of the branch tips on a phylogenetc tree (i.e., which lineage goes to the right and which goes to the left), this ordering is not meaningful at all. Instead, the key to understanding genetic relationships in phylogenetic trees is common ancestry. Common ancestry refers to the fact that distinct descendent lineages have the same ancestral lineage in common with one another, as shown in illustration five.

Determining the dates of tMRCA for Y-DNA haplogroups involves several steps and assumptions, which also come with certain limitations. While SNP-based calculations provide a powerful tool for estimating tMRCA dates, they are subject to limitations related to mutation rate variability, data quality, and the assumptions underlying the models used to estimated their respective dates.

Illustration Five: the Most Recent Common Ancestor

Variability of tMRCA Estimates

Current calculations for TMRCA in Y-DNA phylogenetic trees rely on counting genetic mutations (SNPs and STRs), using probabilistic models that integrate multiple data types, and adjusting results based on historical context and demographic factors. [36]

As with any historical calculations, there are a number of inherent limitations associated with the estimation process. The mutation rate is not perfectly uniform and can vary between different parts of the Y chromosome. This variability can lead to inaccuracies in MRCA date estimates. [37] Random mutations can skew results, especially when comparing individual Big Y results. Anomalies in variant counts can lead to discrepancies in estimated dates. [38]

The calculations rely on assumptions about mutation rates and the models used. Different models or assumptions can yield different estimates, and there is ongoing debate about the most accurate methods. [39] Historical events like bottlenecks or gene flow can affect the genetic diversity of Y-DNA haplogroups, potentially altering the apparent MRCA date. [40]

An example of the variability associated with establishing estimated dates for MRCAs is provided below. Illustration six depicts an high level phylogenetic tree that covers part of my Y-DNA ancestral genetic path. Some of my intermediate MRCAs are not shown in the tree. The tree starts with haplogroup G-L140. The shaded arrow in the illustration depicts the path of my YDNA genetic mutations from haplogroup G-L140 to haplogrop G-Y8903.

Illustration Six: A Philogenetic Tree of haplgroup G2a-L140

Based on the genetic path of haplogroup group mutations shown in the phylogenetic tree, I have chosen four MRCAs shown in table one. The table provides an estimated birth date of each of the MRCAs associated with the unique Y-DNA mutations. Based on the calucations used by FamilyTreeDNA, the table also provides statistical confidence ranges or intervals of the 99, 95 and 68 percent likelihood of the birth dates to fall within a given time range.

Table One: Selected Most Recent Common Ancestors and Estimated Births

MRCA Estimated Birth (Mean)	Estimated Birth Date	99 % Confidence Interval (CI) of when MRCA was born (Calendar Date)	95 % CI Calendar Date Range	65 % CI Calendar Date Range
L140	4,587 BCE	3615 – 1650 BCE	3256 – 1958 BCE	2913 – 2255 BCE
L497	7,549 BCE	7220 – 4051 BCE	6642 – 4549 BCE	6090 – 5028 BCE
Z1817	5,133 BCE	4279 – 2094 BCE	3880 – 2437 BCE	3499 – 2766 BCE
Y8903 / FGC477	4,279 BCE	3374 – 1307 BCE	2989 – 1625 BCE	2624 – 1933 BCE

Source: Scientific Details for Selected FamilyTreeDNA Haplogroups, 8 Mar 2025, FamilyTreeDNA Discover Reports

The wide variations associated with each estimate of birth for the MRCAs underscore the wide variation of age estimates.

A graphic portrayal of the confidence intervals for estimating the birthdate for the MRCA associated with the G-L497 haplogroup is provided in illustration eight. The common ancestor associated with G-L497 is likely to have been born around the year 5524 BCE, but there is a significant range of his estimated birth. There is a 99 percent change that this person could have been born anywhere between around 7220 BCE and 4051 BCE, a variance of 3,169 years. Narrower bands of probability of when this person was born are provided for 95 percent and 68 percent chances.

Illustration Eight: Confidence Interval Ranges for Estimating Birth Date for MRCA for Haplogroup G-L497

What Do Patterns of Subclades in Phylogenetic Trees Tell Us

Different haplogroup clades or sub-branches within the Y-chromosome phylogeneic trees show distinct patterns. The G haplogroup has experienced both the expansion and contraction of subclades through its westward European migratory path.

Illustration Nine

A haplotree with many subclades occurring in a short time period typically indicates a period of rapid population growth. When a Y-DNA phylogenetic tree displays numerous subclades emerging within a short timeframe, this pattern reveals important insights about our ancestral history. This phenomenon, known as a “rapid radiation” or “burst” of lineages, represents a significant demographic event that can tell us much about historical population dynamics and human migrations.

Illustration nine provides an example of this expansion in an E haplogroup branch.

These rapid diversification events often coincide with favorable historical conditions that supported population growth, such as:

Technological innovations that improved survival rates;
Expansion into new, resource-rich territories;
Climate changes that created more favorable living conditions;
Periods of relative peace and prosperity;
Agricultural developments supporting larger populations; and
Many rapid subclade formations correlate with important cultural transitions, such as the adoption of agriculture, metallurgy, or other technological advances that enabled population growth.

The biological mechanism behind rapid subclade formation involves multiple male lineages successfully reproducing around the same time period. Since Y-DNA mutations occur at relatively slow rates, a cluster of branches occurring closely together in evolutionary time suggests numerous male lineages were simultaneously successful in passing on their Y chromosomes. [41]

Typically, approximately every third or fourth generation, a son is born with a SNP that makes him unique and slightly different from his father”. When many such lineages survive in a short time period, it creates a characteristic ‘star-like pattern’ in the phylogenetic tree, with numerous branches emanating from a single ancestral node or MRCA.

This pattern creates an imbalance where larger ‘child’ clades or haplogroup branches receive statistically more mutations than smaller child clades. The mutations occurring early in the expansion become defining features of the larger subsequent subclades. [42]

This clustering of subclades in time can sometimes cause statistical challenges in dating the exact age of these closely-spaced subclades as there may be too few mutations separating parent clades from child clades to establish precise timing of the most recent common ancestor.

This statistical artifact of clustering subclades is evident when looking at the Griff(is)(es)(ith) family lineage in Table Two below. I have noted this by annotating the time passed between subclades in red.

Illustration Ten

Periods of rapid subclade formation stand in stark contrast to periods of slower diversification. When a phylogenetic tree shows a long branch with many accumulated mutations before diversification occurs, this suggests a lineage survived through challenging conditions before eventually flourishing. When a Y-DNA phylogenetic tree displays few subclades over a long stretch of time, this pattern represents what geneticists call a “long branch” – a significant period where little apparent diversification occurred in the paternal lineage. This phenomenon has several important biological, demographic, and methodological implications.

A haplotree with few subclades is provided in illustration ten. Haplogroup E-Y19508 a major branch that has the same most recent common ancestor that is associated with the branch in E-Z5017 in illustration nine. However, the phylogenetic tree associated with the E-Y19508 branch is long and narrow. This is an example of an E haplogroup branch spread over a long time period. This typically indicates slower population growth and more stable demographic conditions.

A primary explanation for long branches with minimal subclade formation is a severe reduction in male effective population size. Studies have documented a pronounced decline in male effective population sizes worldwide around 3000-5000 years ago that was not observed in female lineages. This genetic bottleneck would naturally result in the elimination of many Y-chromosome lineages, leaving fewer surviving male lines to develop subclades. [43]

Geographic isolation and natural barriers can contribute to this pattern by creating separate, isolated populations with limited genetic exchange. The slow accumulation of branches can also result from limited population growth, reduced genetic diversity, or selective pressures affecting Y-chromosome variation. [44]

Long branches with few subclades may also reflect cultural practices that influenced male reproductive success. 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. [45] 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 Y lineages. [46]

When interpreting long branches in the Y-DNA tree, several technical factors must be considered. Long branches may be dueto sampling limitations. Current phylogenetic trees are based on available samples which may not represent all historical populations. For example, the R haplogroup shows 16 times more branching than the G haplogroup despite G being almost twice as old. This could be partly due to sampling biases in European populations. [47] There is a phylogenetic artifact, long branch attraction, where distantly related lineages with significant accumulated changes (YDNA variant mutations) appear to be closely related when they are not. This can create false relationships in analyses of long branches. [48]

Some branches of its subclades have long branches and deep-rooting nodes (ancestors). This is reflected in in two notable historic periods that are associated with my Y-DNA lineage, such as the G-PF3345 and G-FGC7515 haplogroups (see illustration elevin).

Illustration Elevin: G Haplogroups with Long Branches

The expansion and contraction of Y-chromosomal subclades across Europe reflect a complex interplay of demographic migrations, cultural transitions, and genetic drift. Over millennia, paternal lineages associated with haplogroups such as G2a, R1b, R1a, I2a, and N1c1 underwent rapid geographical expansion due to founder effects, male-mediated population movements, and technological innovations. These expansions were often tied to transformative periods in European prehistory, including post-glacial recolonization, the Neolithic Revolution, and Bronze Age pastoralist migrations.

Phylogenetic Comparisons Between European Haplogroups

Phylogenetic resolution refers to how accurately and specifically a phylogenetic tree depicts the evolutionary relationships between tMRCAs. A ‘fully resolved’ tree shows clear, bifurcating relationships with each internal node (most recent common ancestor) having two descendants, while a tree with polytomies (multiple branches emerging from a single node) indicates unresolved relationships. [49]

The phylogenetic resolution of haplogroup G is relatively limited compared to other major European Y-DNA haplogroups, such as haplogroups I and R1a, primarily due to differences in demographic history, geographic dispersal patterns, and population dynamics. Haplogroup G had fewer subclades and limited branching, localized pockets of distribution, strong founder effects and limited genetic diversity, and cultural isolation or assmilation into other cultures through time.

Table Two: Comparison of Phylogenetic Characeristics between Haplogroups G, I and R1a

Aspect	Haplogroup G	Haplogroup I	Haplogroup R1a
Phylogenetic Resolution	Moderate to low; fewer subclades identified, limited branching complexity [50]	High; clearly defined subclades with distinct geographic distributions [51]	High; extensive branching and detailed substructure characterized [52]
Geographic Distribution	Localized pockets (e.g., Alps, Sardinia, Crete); isolated populations with limited gene flow [53]	Widespread across Europe, multiple geographically distinct subclades (e.g., Scandinavia vs. Balkins). [54]	Widely dispersed across Europe and Asia; clear regional substructure (e.g., Z280 in Europe, Z93 in Asia) [55]
Founder Effects / Bottlenecks	Strong founder effects due to Neolithic agricultural expansions from Near East into Europe; limited initial genetic diversity carried forward [56]	Postglacial recolonization from multiple refuge areas; distinct expansions from diverse source populations. [57]	Multiple expansions from Near East/Central Asia; diversification events well-documented through ancient migrations. [58]
Geographic Distribution	Concentrated pockets e.g., Tyrol, Sardinia, Crete; limited clinal patterns; indicative of isolation by distance. [59]	Clear geographic gradients and distinct regional peaks (Scandinavia, Dinaric Alps); clinal patterns evident. [60]	Extensive geographic distribution with clear regional differentiation; basal branches found primarily in Iran/Turkey region. [61]
Cultural/ Demographic Factors	Strongly associated with early Neolithic agricultural expansions; founder effects and cultural isolation restricted diversification. [62]	Associated with postglacial recolonization events and subsequent demographic expansions; multiple regional founder effects created distinct branches. [63]	Associated with Bronze Age Indo-European migrations; rapid expansions from small founder populations allowed clear substructure development [64] .

Limitations Associated with the Use and Interpretation of Y-DNA Phylogenetic Trees

The reconstruction of Y-DNA phylogenetic trees has revolutionized our understanding of paternal lineage evolution, population migrations, and historical demographic processes. However, these analyses are constrained by several technical, methodological, and biological limitations. Y-DNA phylogenies must be interpreted with caution, acknowledging their inherent uncertainties and contextualizing findings within broader genomic and historical frameworks.

Key challenges include variability in mutation rates across haplogroups, biases in sequencing and sampling, limitations of analytical models, and the inherent complexities of the Y chromosome’s non-recombining structure. Additionally, factors such as homoplasy in short tandem repeats (STRs) [65] , evolving nomenclature systems, and population-specific historical events further complicate the interpretation of Y-DNA phylogenies.

A foundational assumption in Y-DNA phylogenetic dating is that mutation rates remain constant across lineages. However, empirical evidence demonstrates significant inter-haplogroup variation in mutation rates. For instance, studies analyzing whole-genome sequences from over 1,700 males revealed up to an 83 percent difference in somatic mutation rates between haplogroups, correlating with phylogenetic branch length heterogeneity. [66] These discrepancies distort time to most recent common ancestor (TMRCA) estimates, as branches with slower mutation rates appear artificially elongated, while rapidly mutating lineages seem younger than their true age. [67]

The reliance on “evolutionary rates” derived from population data or pedigree studies introduces additional uncertainty. [68] This is exacerbated by the tendency of certain STRs to undergo backmutations, which obscure true phylogenetic relationships and inflate TMRCA estimates. [69]

Most Y-DNA data derive from modern populations, with limited ancient DNA representation. This temporal gap complicates efforts to resolve historical migration events or validate putative branching orders. For example, the coalescence time of R1a-M417 (approiximately 5,800 years ago) relies heavily on modern sequences, which may not capture extinct subclades that diversified during the Neolithic or Bronze Age. [70] This may be the case with many of the haplogrous associated with the Griff(is)(es)(ith) patrlineal genetic line.

Source:

Feature Banner: The banner at the top of the story is a portrayal of two phylogenetic trees that depict portions of the G haplogroup migratory route for my terminal haplogroup in Wales.

The phylogenetic tree on the left hand side reflects the phylogenetic tree of Haplogroup G2a-L140. The haplgroup G2a-L140 is most commonly found in Europe, particularly in northern and western regions. The haplogroup is believed to have entered Europe during the Neolithic period, associated with the spread of agriculture. The upstream mutations include M201 > L89 > P15 > L1259 > L30 > L141 > P303 > L140. See Hay, Maciamo, Phylogeny of G2a, Haplogroup G2a, July 2023, Eupedia, https://www.eupedia.com/europe/Haplogroup_G2a_Y-DNA.shtml

The hylogenetic tree on the right is a continuation of the haplogroups linked from one of the common ancestors associated with haplogroup G-Y8903 / FGC477 that is indicated in the phylogenetic tree on the left. The descendant asociated with this haplogroup was born around 2250 BCE. The tree on the right is based on YDNA FamilyTreeDNA test kit results. Names that appear on this chart indicate persons whose YDNA testing results identified a new branch. These SNP branches are hundreds or thousands of years old and each may include many other surnames besides those shown in the chart. Source: Rolf Langland and Mauricio Catelli, Haplogroup G –L497 Chart D: FGC477 Branch, 30 Jan 2025, https://drive.google.com/file/d/1iizSCGkw_8x2cAqm2Evv-b_ZSxY40E1j/view

It is noted that my Y-700 DNA test results identified a new branch, as reflected in the phylogenetic tree. Click here for larger version of the banner image

[1] Zou Y, Zhang Z, Zeng Y, Hu H, Hao Y, Huang S, Li B. Common Methods for Phylogenetic Tree Construction and Their Implementation in R. Bioengineering (Basel). 2024 May 11;11(5):480. doi: 10.3390/bioengineering11050480. PMID: 38790347; PMCID: PMC11117635, https://pmc.ncbi.nlm.nih.gov/articles/PMC11117635/

Understanding phylogenies, Understanding Evolution, Evolution 101, University of California Berkley https://evolution.berkeley.edu/evolution-101/the-history-of-life-looking-at-the-patterns/understanding-phylogenies/

Phylogenetic tree, Wikipedia, This page was last edited on 26 February 2025, https://en.wikipedia.org/wiki/Phylogenetic_tree

Boudreau, Sarah, What’s the difference between a cladogram and a phylogenetic tree?, 28 Apr 2023, Visible Body, https://www.visiblebody.com/blog/phylogenetic-trees-cladograms-and-how-to-read-them

[2] Big Y-700: The Forefront of y Chromosome, 7 Jun 2019, FamilyTreeDNA Blog, https://blog.familytreedna.com/human-y-chromosome-testing-milestones/

Caleb Davis, Michael Sager, Göran Runfeldt, Elliott Greenspan, Arjan Bormans, Bennett Greenspan, and Connie Bormans, Big Y-700 White Paper, 22 Mar 2019, https://blog.familytreedna.com/wp-content/uploads/2019/03/big-y-700-white-paper_compressed.pdf

[3] A most recent common ancestor (MRCA) is the closest individual from whom all members of a specified group of people are directly descended. In genetic genealogy, this concept applies to both biological organisms and groups of genes (haplotypes).

Estes, Roberta, What Does MCRA (MRCA) Really Mean??, 6 Aug 2012, DNAeXplained – Genetic Genealogy, https://dna-explained.com/2012/08/06/what-does-mcra-really-mean/

Most recent common ancestor, International Society of Genetic Genealogy Wiki, This page was last edited on 31 January 2017,https://isogg.org/wiki/Most_recent_common_ancestor

Most common recent ancestor, Wikipedia, This page was last edited on 12 February 2025, https://en.wikipedia.org/wiki/Most_recent_common_ancestor

[4] Spencer, Rob, Data Source and SNP Dates, Discussion, SNP Tracker, https://scaledinnovation.com/gg/snpTracker.html

Batini, C., Hallast, P., Zadik, D., Maisano Delser, P., Benazzo, A., Ghirotto, S., Arroyo-Pardo, E., Cavalleri, G.L., de Knijff, P., Myhre Dupuy, B., Eriksen, H.A, King, T.E., López de Munain, A., López-Parra, A.M., Loutradis, A., Milasin, J., Novelletto, A., Pamjav, H., Sajantila, A., Tolun, A., Winney, B., and JOBLING, M.A. (2015) Large-scale recent expansion of European patrilineages shown by population resequencing. Nature Comm., 6, 7152. doi:10.1038/ncomms8152, (PubMed) https://pubmed.ncbi.nlm.nih.gov/25988751/

Hallast, P., Batini, C., Zadik, D., Maisano Delser, P., Wetton, J.H., Arroyo-Pardo, E., Cavalleri, G.L., de Knijff, P., Destro Bisol, G., Myhre Dupuy, B., Eriksen, H.A, Jorde, L.B., King, T.E., Larmuseau, M.H., López de Munain, A., López-Parra, A.M., Loutradis, A., Milasin, J., Novelletto, A., Pamjav, H., Sajantila, A., Schempp, W., Sears, M., Tolun, A., Tyler-Smith, Van Geystelen, A., Watkins, S., Winney, B., and JOBLING, M.A. (2015) The Y-chromosome tree bursts into leaf: 13,000 high-confidence SNPs covering the majority of known clades. Mol. Biol. Evol., 32, 661–673. doi: 10.1093/molbev/msu327 , (PubMed). https://pubmed.ncbi.nlm.nih.gov/25988751/

Zeng, T.C., Aw, A.J. and Feldman, M.W., 2018. Cultural hitchhiking and competition between patrilineal kin groups explain the post-Neolithic Y-chromosome bottleneck. Nature communications, 9(1), p.2077.

[5] Violatti, Christian Neolithic Period , World History Encyclopedia, 2 Apr 2018 https://www.worldhistory.org/Neolithic/

[6] 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 Commun 9, 2077 (2018), page1, https://doi.org/10.1038/s41467-018-04375-6

[7] Ibid

[8] 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

[9] 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

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

[10] Hay, Maciamo, Phylogenetic trees of Y-chromosomal haplogroups, May 2017, Eupedia, https://www.eupedia.com/genetics/phylogenetic_trees_Y-DNA_haplogroups.shtml

Howard III, William and Frederic R. Schwab, Dating Y-DNA Haplotypes on a Phylogentic Tree: Tying the Genealogy of Pedigrees and Surname Clusters into Genetic Time Scales, Journal of Genetic Genealogy, Volume 7, Number 1 (Fall 2011) Reference Number: 71.005, https://jogg.info/wp-content/uploads/2021/09/71.005.pdf

[11] The animation was produced by a FamilyTreeDNA (FTDNA) online program called Globetrekker ^TM. It is a specialized mapping tool developed by FTDNA as an exclusive feature for their Big Y-DNA test customers. It visualizes ancestral migration paths on a global scale, tracing paternal lineage journeys from “Y-Adam” (the earliest common paternal ancestor, approximately 200,000 years ago) to the most recent known locations of direct paternal ancestors. Globetrekker employs phylogenetic algorithms that factor in geographical topography, historical sea levels, land elevations, and ice age glaciation patterns to determine likely ancestral migration routes.

The following are key features of the Globetrekker program:

Integrated Phylogenetic Tree Browser: An integrated tree browser allows the use to view specific migratory paths based on a chosen terminal haplogroup.

Extensive Data: Globetrekker utilizes the largest Y-DNA tree and a comprehensive database of high-resolution DNA samples, including detailed paternal ancestral information .

Advanced Algorithms: It employs sophisticated phylogenetic algorithms that incorporate topographical data, historical global sea levels, land elevation, and ice age glaciation to accurately reconstruct ancient migration routes .

Historical Maps: The tool provides interactive world maps depicting ancient sea levels and landforms, such as Doggerland during the Last Glacial Maximum .

Personalized Animation: Users receive a customized animation illustrating 200,000 years of their paternal lineage history .

Extensive Migration Paths: Globetrekker currently includes over 48,000 paternal line migration paths covering every populated continent, with new paths regularly added .

Globetrekker’s main limitation is the relatively small number of available Big Y-DNA samples. As more individuals participate in Big Y testing, the accuracy and granularity of migration paths are expected to improve significantly over time. The video is based on the migration mapping for the terminal haplogroup for G-Y132505.

Estes, Roberta, Globetrekker – A New Feature for Big Y Customers from FamilyTreeDNA, 4 Aug 2023, DNAeXplained – Genetic Genealogy, https://dna-explained.com/2023/08/04/globetrekker-a-new-feature-for-big-y-customers-from-familytreedna/

Runfeldt, Goran , Globertrekker, Part 1: A NewFamilyTreeDNA Discover™ Report that Puts Big Y on the Map, 31 Jul 2023, FamilyTreeDNA Blog, https://blog.familytreedna.com/globetrekker-discover-report/

Maier, Paul, Globetrekker, Part 2: Advancing the Science of Phylogeography, 15 Aug 2023, FamilyTreeDNA Blog, https://blog.familytreedna.com/globetrekker-analysis/

Vilar, Miguel, Globetrekker, Part 3: We Are Making History, 26 Sep 2023, FamilyTreeDNA Blog, https://blog.familytreedna.com/globetrekker-history/

[12] Doggerland was a vast landmass that once connected the British Isles to mainland Europe, encompassing areas now submerged beneath the North Sea and the English Channel. Named after Dogger Bank, a submerged sandbank frequented by Dutch fishing vessels known as “doggers,” Doggerland existed primarily during the Late Pleistocene and Early Holocene periods, approximately 10,000 to 6,500 years ago.

Doggerland, Wikipedia, This page was last edited on 10 March 2025, https://en.wikipedia.org/wiki/Doggerland

James Walker, Vincent Gaffney, Simon Fitch, Merle Muru, Andrew Fraser, Martin Bates and Richard Bates, A great wave: the Storegga tsunami and the end of Doggerland?, Antiquity , Volume 94 , Issue 378 , December 2020 , pp. 1409 – 1425 DOI: https://doi.org/10.15184/aqy.2020.49 , https://www.cambridge.org/core/journals/antiquity/article/great-wave-the-storegga-tsunami-and-the-end-of-doggerland/CB2E132445086D868BF508041CC1B827#article

Urbanus, Jason, Mapping a Vanished Landscape, Archaelogy magazine, March/April 2022, https://archaeology.org/issues/march-april-2022/letters-from/doggerland-mesolithic-submerged-landscape/

De Abreu, Kristine, Exploration Mysteries: Doggerland, 13 Feb 2024, Explorersweb, https://explorersweb.com/exploration-mysteries-doggerland/

[13] Balaresque P, Bowden GR, Adams SM, Leung HY, King TE, Rosser ZH, Goodwin J, Moisan JP, Richard C, Millward A, Demaine AG, Barbujani G, Previderè C, Wilson IJ, Tyler-Smith C, Jobling MA. A predominantly neolithic origin for European paternal lineages. PLoS Biol. 2010 Jan 19;8(1):e1000285. doi: 10.1371/journal.pbio.1000285. PMID: 20087410; PMCID: PMC2799514, PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC8228294/

Semino O, Magri C, Benuzzi G, Lin AA, Al-Zahery N, Battaglia V, Maccioni L, Triantaphyllidis C, Shen P, Oefner PJ, Zhivotovsky LA, King R, Torroni A, Cavalli-Sforza LL, Underhill PA, Santachiara-Benerecetti AS. Origin, diffusion, and differentiation of Y-chromosome haplogroups E and J: inferences on the neolithization of Europe and later migratory events in the Mediterranean area. Am J Hum Genet. 2004 May;74(5):1023-34. doi: 10.1086/386295. Epub 2004 Apr 6. PMID: 15069642; PMCID: PMC1181965, (PubMed)https://pmc.ncbi.nlm.nih.gov/articles/PMC1181965

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E.K. Khusnutdinova, N.V. Ekomasova, M.A. Dzhaubermezov, L.R. Gabidullina, Z.R. Sufianova1, I.M. Khidiyatova, A.V. Kazantseva, S.S. Litvinov, A.Kh. Nurgalieva, D.S. Prokofieva, Distribution of Haplogroup G-P15 of the Y-chromosome Among Representatives of Ancient cultures and Modern Populations of Northern Eurasia, Opera Med Physiol. 2023. Vol. 10 (4), 57-72, doi: 10.24412/2500-2295-2023-4-57-72, https://operamedphys.org/system/tdf/pdf/06_DISTRIBUTION%20OF%20HAPLOGROUP%20G-P15_0.pdf?file=1&type=node&id=555&force=0

Maciamo, Hay, Phylogeny of G2a, Haplogroup G2a, July 2023, Eupedia, https://www.eupedia.com/europe/Haplogroup_G2a_Y-DNA.shtml

[14] The Neolithic agricultural expansion, also known as the Neolithic Revolution, was a pivotal period in human history marked by the transition from hunter-gatherer lifestyles to settled agricultural communities, starting around 10,000 years ago. Agricultural and husbandry practices originated 10,000 years ago in a region of the Near East known as the Fertile Crescent. According to the archaeological record this phenomenon, known as “Neolithic”, rapidly expanded from these territories into Europe.

Main Archaeological Sites of the Pre-Pottery Neolithic period, BCE c. 7500, in the “Fertile Crescent”

Source: Neolithic Revolution, Wikipedia, This page was last edited on 1 March 2025, https://en.wikipedia.org/wiki/Neolithic_Revolution

Mesolithic Tribes and the Origins of Agriculture in the Near East (9000-7000 BCE)

[15] Neolithic, Wikipedia, This page was last edited on 18 March 2025, https://en.wikipedia.org/wiki/Neolithic

[16] Ancient DNA from the Treilles group in southern France (c. 3000 BCE) revealed that nintey percent of male remains belonged to G2a, with mitochondrial DNA (mtDNA) showing affinity to Neolithic Aegean populations . This genetic profile supports a maritime migration route linking Anatolia to Iberia via Crete and the Adriatic. Notably, the absence of the N1a mtDNA haplogroup—common in Central European Neolithic groups—in Treilles samples underscores the genetic distinctiveness of Mediterranean versus Danubian Neolithic expansions .

M. Lacan, C. Keyser, F. Ricaut, N. Brucato, F. Duranthon, J. Guilaine, E. Crubézy, & B. Ludes, Ancient DNA reveals male diffusion through the Neolithic Mediterranean route, Proc. Natl. Acad. Sci. U.S.A. 108 (24) 9788-9791,https://doi.org/10.1073/pnas.1100723108 (2011)

Anna Szécsényi-Nagy , Guido Brandt , Wolfgang Haak , Victoria Keerl , János Jakucs , Sabine Möller-Rieker , Kitti Köhler , Balázs Gusztáv Mende , Krisztián Oross , Tibor Marton , Anett Osztás , Viktória Kiss , Marc Fecher , György Pálfi , Erika Molnár , et al, Tracing the genetic origin of Europe’s first farmers reveals insights into their social organization, 22 Apr 2015, Volume 282, Issue 1805, Proceedings of the royal Society Biological Sciences, https://royalsocietypublishing.org/doi/10.1098/rspb.2015.0339

[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

Anna Szécsényi-Nagy , Guido Brandt , Wolfgang Haak , Victoria Keerl , János Jakucs , Sabine Möller-Rieker , Kitti Köhler , Balázs Gusztáv Mende , Krisztián Oross , Tibor Marton , Anett Osztás , Viktória Kiss , Marc Fecher , György Pálfi , Erika Molnár , et al, Tracing the genetic origin of Europe’s first farmers reveals insights into their social organization, 22 Apr 2015, Volume 282, Issue 1805, Proceedings of the royal Society Biological Sciences, https://royalsocietypublishing.org/doi/10.1098/rspb.2015.0339

Myres NM, Rootsi S, Lin AA, Järve M, King RJ, Kutuev I, Cabrera VM, Khusnutdinova EK, Pshenichnov A, Yunusbayev B, Balanovsky O, Balanovska E, Rudan P, Baldovic M, Herrera RJ, Chiaroni J, Di Cristofaro J, Villems R, Kivisild T, Underhill PA. A major Y-chromosome haplogroup R1b Holocene era founder effect in Central and Western Europe. Eur J Hum Genet. 2011 Jan;19(1):95-101. doi: 10.1038/ejhg.2010.146. Epub 2010 Aug 25. PMID: 20736979; PMCID: PMC3039512, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC3039512/

[18] 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/

[19] Myres NM, Rootsi S, Lin AA, Järve M, King RJ, Kutuev I, Cabrera VM, Khusnutdinova EK, Pshenichnov A, Yunusbayev B, Balanovsky O, Balanovska E, Rudan P, Baldovic M, Herrera RJ, Chiaroni J, Di Cristofaro J, Villems R, Kivisild T, Underhill PA. A major Y-chromosome haplogroup R1b Holocene era founder effect in Central and Western Europe. Eur J Hum Genet. 2011 Jan;19(1):95-101. doi: 10.1038/ejhg.2010.146. Epub 2010 Aug 25. PMID: 20736979; PMCID: PMC3039512, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC3039512/

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

[20] Chiaroni J, Underhill PA, Cavalli-Sforza LL. Y chromosome diversity, human expansion, drift, and cultural evolution. Proc Natl Acad Sci U S A. 2009 Dec 1;106(48):20174-9. doi: 10.1073/pnas.0910803106. Epub 2009 Nov 17. Erratum in: Proc Natl Acad Sci U S A. 2010 Jul 27;107(30):13556. PMID: 19920170; PMCID: PMC2787129, PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC2787129/

Myres NM, Rootsi S, Lin AA, et al, A major Y-chromosome haplogroup R1b Holocene era founder effect in Central and Western Europe. Eur J Hum Genet. 2011 Jan;19(1):95-101. doi: 10.1038/ejhg.2010.146. Epub 2010 Aug 25. PMID: 20736979; PMCID: PMC3039512, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC3039512/

[21] Penske, S., Rohrlach, A.B., Childebayeva, A. et al. Early contact between late farming and pastoralist societies in southeastern Europe. Nature 620, 358–365 (2023). https://doi.org/10.1038/s41586-023-06334-8

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

Lamnidis, T.C., Majander, K., Jeong, C. et al. Ancient Fennoscandian genomes reveal origin and spread of Siberian ancestry in Europe. Nat Commun 9, 5018 (2018). https://doi.org/10.1038/s41467-018-07483-5

[23] Hay, Maciamo, Phylogenetic trees of Y-chromosomal haplogroups, May 2017, Eupedia, https://www.eupedia.com/genetics/phylogenetic_trees_Y-DNA_haplogroups.shtml#IntroductionHuman Y-chromosome DNA haplogroup, Wikipedia, This page was last edited on 31 December 2024, https://en.wikipedia.org/wiki/Human_Y-chromosome_DNA_haplogroup

Dunn, Casey W., Chapter 9 Phylogenies and time, Phylogenetic Biology, 28 Oct 2024, Text for course, Phylogenetic Biology (Yale EEB354), licensed under the Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License. It is available to read online for free at https://dunnlab.org/phylogenetic_biology/phylogenies.html#trees-branch-lengths

Hay, Maciamo, Phylogenetic trees of Y-chromosomal haplogroups, May 2017, Eupedia, https://www.eupedia.com/genetics/phylogenetic_trees_Y-DNA_haplogroups.shtml#Introduction

[24] 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

[25] Hay, Maciamo, Phylogenetic trees of Y-chromosomal haplogroups, May 2017, Eupedia, https://www.eupedia.com/genetics/phylogenetic_trees_Y-DNA_haplogroups.shtml#IntroductionHuman Y-chromosome DNA haplogroup, Wikipedia, This page was last edited on 31 December 2024, https://en.wikipedia.org/wiki/Human_Y-chromosome_DNA_haplogroup

[26] These estimates derive from large-scale sequencing datasets, pedigree studies, and comparative analyses of haplogroup differentiations. Key factors influencing this range include the coverage of the male-specific Y chromosome (MSY) region, the mutation rate per base pair, and the statistical models used to account for uncertainties in SNP counting and temporal calibration. [26a]

Mutation rate estimates differ across sequencing technologies. There are three notable testing platforms. The FamilyTreeDNA (FTDNA) Big Y-700 test analyzes approximately 14.6 million base pairs, yielding an average mutation rate of 83–85 years per SNP. This estimate, derived from YDNA Warehouse data, reflects high-coverage regions deemed reliable for genealogical applications. [26b] The FTDNA BigY-500 test covers 9.3 million base pairs, resulting in a slower rate of 131 years per SNP due to reduced coverage compared to BigY-700. [26c] The YFull (ComBed) coverage test uses 8.5 million base pairs and reports 144 years per SNP, prioritizing conservative regions (comBED) to minimize false positives. [26d]

Based on academic and ‘consensus’ estimates, evolutionary rates, calibrated using ancient DNA or historical events, suggest 0.75–0.89 substitutions per billion base pairs per year (equivalent to 83–89 years/SNP for typical sequencing lengths). Genealogical (pedigree) rates, observed in father-son studies, are slightly faster due to shorter generational intervals. Iain McDonald’s analysis of 15 million base pairs estimates 83–186 years per SNP, with higher values reflecting conservative adjustments for regions with variable coverage. [26e]

[26a] Irvine, James M., Y-DNA SNP-Based TMRCA Calculations for Surname Project Administrators, Journal of Genetic Genealogy, Volume 9, Number 1 (Fall 2021), Reference Number: 91.007, https://jogg.info/wp-content/uploads/2021/12/91.007-Article.pdf

SNP Dating, Genomic Genealogy Research, University of Strathclyde Glasgow, https://www.strath.ac.uk/studywithus/centreforlifelonglearning/genealogy/geneticgenealogyresearch/snpdating/

Balanovsky O. Toward a consensus on SNP and STR mutation rates on the human Y-chromosome. Hum Genet. 2017 May;136(5):575-590. doi: 10.1007/s00439-017-1805-8. Epub 2017 Apr 28. PMID: 28455625, (PubMed) https://pubmed.ncbi.nlm.nih.gov/28455625/

[26b] SNP Dating, Genomic Genealogy Research, University of Strathclyde Glasgow, https://www.strath.ac.uk/studywithus/centreforlifelonglearning/genealogy/geneticgenealogyresearch/snpdating/

[26c] McDonald, Ian, SNP-based age analysis methodology: a summary

Summarised description of the age analysis pipeline, June 2017, https://www.jb.man.ac.uk/~mcdonald/genetics/pipeline-summary.pdf

[26d] Ibid

[26e] Balanovsky O. Toward a consensus on SNP and STR mutation rates on the human Y-chromosome. Hum Genet. 2017 May;136(5):575-590. doi: 10.1007/s00439-017-1805-8. Epub 2017 Apr 28. PMID: 28455625, (PubMed) https://pubmed.ncbi.nlm.nih.gov/28455625/

McDonald, Ian, SNP-based age analysis methodology: a summary, Summarised description of the age analysis pipeline, June 2017, https://www.jb.man.ac.uk/~mcdonald/genetics/pipeline-summary.pdf

[27] The interpretation of branch lengths depends heavily on mutation rate calculations. The standard deviation in branch lengths from high-coverage sequences is relatively low (around 4 percent). This allows for precise temporal estimates. High-coverage DNA sequencing has identified mutation rates of approximately 2-3 base pairs per generation.

Jeanson, Nathaniel, 4 Dec, 2019, Answers Research Journal (ARJ), 12: 405-423, https://answersresearchjournal.org/human-y-chromosome-molecular-clock/

There is ongoing disagreement about the temporal interpretation of Y-chromosome branch lengths. Some researchers argue for a longer timescale of 120-156 thousand years to the most recent common ancestor while others propose much shorter timescales of just a few thousand years. See:

Jeanson, Nathaniel, 4 Dec, 2019, Answers Research Journal (ARJ), 12: 405-423, https://answersresearchjournal.org/human-y-chromosome-molecular-clock/

Poznik GD, Henn BM, Yee MC, Sliwerska E, Euskirchen GM, Lin AA, Snyder M, Quintana-Murci L, Kidd JM, Underhill PA, Bustamante CD. Sequencing Y chromosomes resolves discrepancy in time to common ancestor of males versus females. Science. 2013 Aug 2;341(6145):562-5. doi: 10.1126/science.1237619. PMID: 23908239; PMCID: PMC4032117, https://pmc.ncbi.nlm.nih.gov/articles/PMC4032117/

[28] Dunn, Casey W., Chapter 9 Phylogenies and time, Phylogenetic Biology, 28 Oct 2024, Text for course, Phylogenetic Biology (Yale EEB354), licensed under the Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License. It is available to read online for free at http://dunnlab.org/phylogenetic_biology/

[29] Poznik GD, et al, Sequencing Y chromosomes resolves discrepancy in time to common ancestor of males versus females. Science. 2013 Aug 2;341(6145):562-5. doi: 10.1126/science.1237619. PMID: 23908239; PMCID: PMC4032117, https://pmc.ncbi.nlm.nih.gov/articles/PMC4032117/

[30] Cruciani F, Trombetta B, Massaia A, Destro-Bisol G, Sellitto D, Scozzari R. A revised root for the human Y chromosomal phylogenetic tree: the origin of patrilineal diversity in Africa. Am J Hum Genet. 2011 Jun 10;88(6):814-818. doi: 10.1016/j.ajhg.2011.05.002. Epub 2011 May 27. PMID: 21601174; PMCID: PMC3113241, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC3113241/

[31] Y Chromosome Consortium. A nomenclature system for the tree of human Y-chromosomal binary haplogroups. Genome Res. 2002 Feb;12(2):339-48. doi: 10.1101/gr.217602. PMID: 11827954; PMCID: PMC155271, 9PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC155271/

Human Y-chromosome DNA haplogroup, Wikipedia, This page was last edited on 31 December 2024, https://en.wikipedia.org/wiki/Human_Y-chromosome_DNA_haplogroup

[32] Hay, Maciamo, Phylogenetic trees of Y-chromosomal haplogroups, May 2017, Eupedia, https://www.eupedia.com/genetics/phylogenetic_trees_Y-DNA_haplogroups.shtml#Introduction

[33] Tunde I. Huszar, Mark A. Jobling, Jon H. Wetton,
A phylogenetic framework facilitates Y-STR variant discovery and classification via massively parallel sequencing, Forensic Science International: Genetics, Volume 35, 2018, Pages 97-106, ISSN 1872-4973, https://doi.org/10.1016/j.fsigen.2018.03.012.
(https://www.sciencedirect.com/science/article/pii/S1872497318300279 )

Phylogenetics, Wikipedia, This page was last edited on 12 February 2025, https://en.wikipedia.org/wiki/Phylogenetics

[34] Rowe-Schurwanz, Katy, How Y-DNA Testing Works, 3 Jun 2024, FamilyTreeDNA Blog, https://blog.familytreedna.com/how-y-dna-testing-works/

Y chromosome DNA tests, International Society of Genetic Genealogy Wiki, This page was last edited on 6 September 2024, https://isogg.org/wiki/Y_chromosome_DNA_tests

[35] William E. Howard III and Frederic R. Schwab, Dating Y-DNA Haplotypes on a Phylogenetic Tree: Tying the Genealogy of Pedgrees and Surname Clusters into Genetic Time Scales, Journal of Genetic Genealogy, Volume 7, Number 1 (Fall 2011) Reference Number: 71.005, https://jogg.info/wp-content/uploads/2021/09/71.005.pdf

Köksal Z, Børsting C, Gusmão L, Pereira V. SNPtotree-Resolving the Phylogeny of SNPs on Non-Recombining DNA. Genes (Basel). 2023 Sep 22;14(10):1837. doi: 10.3390/genes14101837. PMID: 37895186; PMCID: PMC10606150, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC10606150/

Tunde I. Huszar, Mark A. Jobling, Jon H. Wetton, A phylogenetic framework facilitates Y-STR variant discovery and classification via massively parallel sequencing, Forensic Science International: Genetics, Volume 35, 2018, Pages 97-106, ISSN 1872-4973,
https://doi.org/10.1016/j.fsigen.2018.03.012

Hay, Maciamo, Phylogenetic trees of Y-chromosomal haplogroups, May 2017, Eupedia, https://www.eupedia.com/genetics/phylogenetic_trees_Y-DNA_haplogroups.shtml

[36] Determination of MRCA Dates”

Calculation Models: The coalescence age (time to MRCA) is calculated using probabilistic models that consider the number of mutations and the mutation rate. These models can be refined with more data and improved algorithms 1 4.

Mutation Rate: The process relies on the concept of a’ molecular clock’, which assumes that mutations occur at a relatively constant rate over time. This rate is typically measured in mutations per base pair per year. For Y-DNA, mutations are often counted as Single Nucleotide Polymorphisms (SNPs) 1 2.

SNP Counting: Full Y-DNA sequencing tests, such as those from Full Genomes Corp. or FamilyTreeDNA’s Big Y, identify novel SNPs. The number of these SNPs, combined with the mutation rate, helps estimate the time to the MRCA. Different tests may yield different mutation rates; for example, Full Genomes Corp. suggests a mutation every 88 years, while Big Y suggests one every 150 years2.

McDonald Ian. Improved Models of Coalescence Ages of Y-DNA Haplogroups. Genes (Basel). 2021 Jun 4;12(6):862. doi: 10.3390/genes12060862. PMID: 34200049; PMCID: PMC8228294, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC8228294/

Irvine, James M., Y-DNA SNP-Based TMRCA Calculations for Surname Project Administrators, Journal of Genetic Genealogy, Volume 9, Number 1 (Fall 2021), Reference Number: 91.007, https://jogg.info/wp-content/uploads/2021/12/91.007-Article.pdf

FamilyTreeDNA Enhances TMRCA Estimates for Improved Family History Research, 9 Sep 2022, FamilyTreeDNA Blog, https://blog.familytreedna.com/tmrca-age-estimates-update/

Walsh B. Estimating the time to the most recent common ancestor for the Y chromosome or mitochondrial DNA for a pair of individuals. Genetics. 2001 Jun;158(2):897-912. doi: 10.1093/genetics/158.2.897. PMID: 11404350; PMCID: PMC1461668, https://pmc.ncbi.nlm.nih.gov/articles/PMC1461668/

Cummings, Karen, Y-DNA: New Tools from FamilyTreeDNA, Professional family History, https://www.professionalfamilyhistory.co.uk/blog/new-y-dna-new-tools-from-familytreedna

Estes, Roberta, The Big Y-700 Test Marries Science to Genealogy, 11 Jul 2024, DNAeXplained – Genetic Genealology, https://dna-explained.com/category/mrca-most-recent-common-ancestor/

Karmin M, Saag L, Vicente M, Wilson Sayres MA, et all 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, https://pmc.ncbi.nlm.nih.gov/articles/PMC4381518/

Bruce Walsh, Estimating the Time to the Most Recent Common Ancestor for the Y chromosome or Mitochondrial DNA for a Pair of Individuals, Genetics, Volume 158, Issue 2, 1 June 2001, Pages 897–912, https://doi.org/10.1093/genetics/158.2.897

Qian, X., Hou, J., Wang, Z. et al. Next Generation Sequencing Plus (NGS+) with Y-chromosomal Markers for Forensic Pedigree Searches. Sci Rep 7, 11324 (2017). https://doi.org/10.1038/s41598-017-11955-x

[37] McDonald Ian. Improved Models of Coalescence Ages of Y-DNA Haplogroups. Genes (Basel). 2021 Jun 4;12(6):862. doi: 10.3390/genes12060862. PMID: 34200049; PMCID: PMC8228294, (PubMed) https://pmc.ncbi.nlm.nih.gov/articles/PMC8228294/

[38] McDonald Ian. Improved Models of Coalescence Ages of Y-DNA Haplogroups.

[39] Poznik GD, Henn BM, Yee MC, Sliwerska E, Euskirchen GM, Lin AA, Snyder M, Quintana-Murci L, Kidd JM, Underhill PA, Bustamante CD. Sequencing Y chromosomes resolves discrepancy in time to common ancestor of males versus females. Science. 2013 Aug 2;341(6145):562-5. doi: 10.1126/science.1237619. PMID: 23908239; PMCID: PMC4032117, https://pmc.ncbi.nlm.nih.gov/articles/PMC4032117/

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