archaeogenetics Archaeogenetics is the study of ancient DNA using various molecular genetic methods and DNA resources. This form of genetic analysis can be applied to human, animal, and plant specimens. Ancient DNA can be extracted from various fossilized specimen ...

, the term Eastern Hunter-Gatherer (EHG), sometimes East European Hunter-Gatherer, or Eastern European Hunter-Gatherer is the name given to a distinct ancestral component that represents descent from

Mesolithic The Mesolithic ( Greek: μέσος, ''mesos'' 'middle' + λίθος, ''lithos'' 'stone') or Middle Stone Age is the Old World archaeological period between the Upper Paleolithic and the Neolithic. The term Epipaleolithic is often used synonymo ...

hunter-gatherer A traditional hunter-gatherer or forager is a human living an ancestrally derived lifestyle in which most or all food is obtained by foraging, that is, by gathering food from local sources, especially edible wild plants but also insects, fung ...

s of

Eastern Europe Eastern Europe is a subregion of the European continent. As a largely ambiguous term, it has a wide range of geopolitical, geographical, ethnic, cultural, and socio-economic connotations. The vast majority of the region is covered by Russia, whi ...

. During the Mesolithic, the EHGs inhabited an area stretching from the

Baltic Sea The Baltic Sea is an arm of the Atlantic Ocean that is enclosed by Denmark, Estonia, Finland, Germany, Latvia, Lithuania, Poland, Russia, Sweden and the North and Central European Plain. The sea stretches from 53°N to 66°N latitude and from ...

to the

Urals The Ural Mountains ( ; rus, Ура́льские го́ры, r=Uralskiye gory, p=ʊˈralʲskʲɪjə ˈɡorɨ; ba, Урал тауҙары) or simply the Urals, are a mountain range that runs approximately from north to south through western ...

and downwards to the

Pontic–Caspian steppe The Pontic–Caspian steppe, formed by the Caspian steppe and the Pontic steppe, is the steppeland stretching from the northern shores of the Black Sea (the Pontus Euxinus of antiquity) to the northern area around the Caspian Sea. It extend ...

. Along with Scandinavian Hunter-Gatherers (SHG) and

Western Hunter-Gatherer In archaeogenetics, the term Western Hunter-Gatherer (WHG), West European Hunter-Gatherer or Western European Hunter-Gatherer names a distinct ancestral component of modern Europeans, representing descent from a population of Mesolithic hunter-gat ...

s (WHG), the EHGs constituted one of the three main genetic groups in the postglacial period of early

Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togeth ...

Europe. The border between WHGs and EHGs ran roughly from the lower

Danube The Danube ( ; ) is a river that was once a long-standing frontier of the Roman Empire and today connects 10 European countries, running through their territories or being a border. Originating in Germany, the Danube flows southeast for , pa ...

, northward along the western forests of the

Dnieper } The Dnieper () or Dnipro (); , ; . is one of the major transboundary rivers of Europe, rising in the Valdai Hills near Smolensk, Russia, before flowing through Belarus and Ukraine to the Black Sea. It is the longest river of Ukraine an ...

towards the western

. According to a study by Wang et al. (2019), EHGs have about 9%

Ancient North Eurasian In archaeogenetics, the term Ancient North Eurasian (generally abbreviated as ANE) is the name given to an ancestral component that represents a lineage ancestral to the people of the Mal'ta–Buret' culture and populations closely related to th ...

(ANE) and 91% descended from a group more closely related to but distinct from WHGs. During the

Neolithic The Neolithic period, or New Stone Age, is an Old World archaeological period and the final division of the Stone Age. It saw the Neolithic Revolution, a wide-ranging set of developments that appear to have arisen independently in several pa ...

and early

Eneolithic The asterisk ( ), from Late Latin , from Ancient Greek , ''asteriskos'', "little star", is a typographical symbol. It is so called because it resembles a conventional image of a heraldic star. Computer scientists and mathematicians often v ...

, likely during the

4th millennium BC The 4th millennium BC spanned the years 4000 BC to 3001 BC. Some of the major changes in human culture during this time included the beginning of the Bronze Age and the invention of writing, which played a major role in starting recorded history. ...

EHGs on the Pontic–Caspian steppe mixed with Caucasus hunter-gatherers (CHGs) with the resulting population, almost half-EHG and half-CHG, forming the

Yamnaya culture The Yamnaya culture or the Yamna culture (russian: Ямная культура, ua, Ямна культура lit. 'culture of pits'), also known as the Pit Grave culture or Ochre Grave culture, was a late Copper Age to early Bronze Age archa ...

. The genetic cluster formed from this admixture is known as

Western Steppe Herder In archaeogenetics, the term Western Steppe Herders (WSH), or Western Steppe Pastoralists, is the name given to a distinct ancestral component first identified in individuals from the Eneolithic steppe around the turn of the 5th millennium BCE, ...

(WSH). Populations closely related to the people of the Yamnaya culture are supposed to have embarked on a massive migration leading to the spread of

Indo-European languages The Indo-European languages are a language family native to the overwhelming majority of Europe, the Iranian plateau, and the northern Indian subcontinent. Some European languages of this family, English, French, Portuguese, Russian, D ...

throughout large parts of Eurasia.

Research

Haak et al. (2015) identified the Eastern Hunter-Gatherers (EHG) as a distinct genetic cluster in two males only. The EHG male of

Samara Samara ( rus, Сама́ра, p=sɐˈmarə), known from 1935 to 1991 as Kuybyshev (; ), is the largest city and administrative centre of Samara Oblast. The city is located at the confluence of the Volga and the Samara rivers, with a population ...

(dated to ca. 5650-5550 BC) carried

Y-haplogroup In human genetics, a human Y-chromosome DNA haplogroup is a haplogroup defined by mutations in the non- recombining portions of DNA from the male-specific Y chromosome (called Y-DNA). Many people within a haplogroup share similar numbers of ...

R1b1a1a* and mt-haplogroup U5a1d. The other EHG male, buried in

Karelia Karelia ( Karelian and fi, Karjala, ; rus, Каре́лия, links=y, r=Karélija, p=kɐˈrʲelʲɪjə, historically ''Korjela''; sv, Karelen), the land of the Karelian people, is an area in Northern Europe of historical significance fo ...

(dated to ca. 5500-5000 BC) carried Y-haplogroup

R1a1 Haplogroup R1a, or haplogroup R-M420, is a human Y-chromosome DNA haplogroup which is distributed in a large region in Eurasia, extending from Scandinavia and Central Europe to southern Siberia and South Asia. While R1a originated c. 22,000 ...

and mt-haplogoup C1g. The authors of the study also identified a

(WHG) cluster and a Scandinavian Hunter-Gatherer (SHG) cluster, intermediate between WHG and EHG. Also Lazaridis et al. (2016) confirmed SHGs to be a mix of EHGs and WHGs. They suggested that EHGs harbored mixed ancestry from

s (ANEs) and WHGs. The people of the

were found to be a mix of EHG and a "Near Eastern related population". During the 3rd millennium BC, the Yamnaya people embarked on a massive expansion throughout

Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...

, which significantly altered the genetic landscape of the continent. The expansion gave rise to cultures such as

Corded Ware The Corded Ware culture comprises a broad archaeological horizon of Europe between ca. 3000 BC – 2350 BC, thus from the late Neolithic, through the Copper Age, and ending in the early Bronze Age. Corded Ware culture encompassed a ...

, and was possibly the source of the distribution of

in Europe. EHGs may have mixed with "an Armenian-like Near Eastern source", which formed the Yamnaya culture, as early as the

(5200-4000 BC). Researchers have found that EHGs may have derived 75% of their ancestry from ANEs. WHGs were in turn a mix of EHGs and the

Upper Paleolithic The Upper Paleolithic (or Upper Palaeolithic) is the third and last subdivision of the Paleolithic or Old Stone Age. Very broadly, it dates to between 50,000 and 12,000 years ago (the beginning of the Holocene), according to some theories coin ...

people (

Cro-Magnon Early European modern humans (EEMH), or Cro-Magnons, were the first early modern humans (''Homo sapiens'') to settle in Europe, migrating from Western Asia, continuously occupying the continent possibly from as early as 56,800 years ago. They i ...

) of the

Grotte du Bichon Grotte du Bichon is a karstic cave in the Swiss Jura, overlooking the river Doubs at an altitude of 846 m, some 5 km north of La Chaux-de-Fonds. It is the site of the discovery of the skeleton a hunter-gatherer of the Azilian (late U ...

Switzerland ). Swiss law does not designate a ''capital'' as such, but the federal parliament and government are installed in Bern, while other federal institutions, such as the federal courts, are in other cities (Bellinzona, Lausanne, Luzern, Neuchâtel ...

. The people of the Mesolithic

Kunda culture The Kunda culture, originating from the Swiderian culture, comprised mesolithic hunter-gatherer communities of the Baltic forest zone extending eastwards through Latvia into northern Russia, dating to the period 8500–5000 BC according to calib ...

and the

Narva culture Narva culture or eastern Baltic was a European Neolithic archaeological culture found in present-day Estonia, Latvia, Lithuania, Kaliningrad Oblast (former East Prussia), and adjacent portions of Poland, Belarus and Russia. A successor of ...

of the eastern

Baltic Baltic may refer to: Peoples and languages *Baltic languages, a subfamily of Indo-European languages, including Lithuanian, Latvian and extinct Old Prussian *Balts (or Baltic peoples), ethnic groups speaking the Baltic languages and/or originatin ...

were a mix of WHG and EHG, showing the closest affinity with WHG. Samples from the

Ukrainian Ukrainian may refer to: * Something of, from, or related to Ukraine * Something relating to Ukrainians, an East Slavic people from Eastern Europe * Something relating to demographics of Ukraine in terms of demography and population of Ukraine * So ...

Mesolithic and

were found to cluster tightly together between WHG and EHG, suggesting genetic continuity in the

Dnieper Rapids The Dnieper Rapids ( uk, Дніпрові пороги, ) are the historical rapids on the Dnieper river in Ukraine, composed of outcrops of granites, gneisses and other types of bedrock of the Ukrainian Shield. The rapids began below the present ...

for a period of 4,000 years. The Ukrainian samples belonged exclusively to the maternal haplogroup U, which is found in around 80% of all European hunter-gatherer samples.

The people of the

Pit–Comb Ware culture The Comb Ceramic culture or Pit-Comb Ware culture, often abbreviated as CCC or PCW, was a northeast European culture characterised by its Pit–Comb Ware. It existed from around 4200 BCE to around 2000 BCE. The bearers of the Comb Ceramic cultu ...

(PCW/CCC) of the eastern Baltic bear 65% EHG ancestry. This is in contrast to earlier hunter-gatherers in the area, who were more closely related to WHG. This was demonstrated using a sample of Y-DNA extracted from a Pit–Comb Ware individual. This belonged to R1a15-YP172. The four samples of mtDNA extracted constituted two samples of U5b1d1, one sample of U5a2d, and one sample of U4a. Günther et al. (2018) analyzed 13 SHGs and found all of them to be of EHG ancestry. Generally, SHGs from western and northern Scandinavia had more EHG ancestry (ca 49%) than individuals from eastern Scandinavia (ca. 38%). The authors suggested that the SHGs were a mix of WHGs who had migrated into Scandinavia from the south, and EHGs who had later migrated into Scandinavia from the northeast along the

Norwegian Norwegian, Norwayan, or Norsk may refer to: *Something of, from, or related to Norway, a country in northwestern Europe * Norwegians, both a nation and an ethnic group native to Norway * Demographics of Norway *The Norwegian language, including ...

coast. SHGs displayed higher frequences of genetic variants that cause light skin ( SLC45A2 and

SLC24A5 SLC may refer to: Places * Salt Lake City, Utah * Salt Lake City International Airport, IATA Airport Code Education * Sarah Lawrence College, NY * School Leaving Certificate (Nepal) * St. Lawrence College, Ontario, Canada * Small Learning Com ...

), and light eyes ( OCA/Herc2), than EHGs and WHGs. Members of the Kunda culture and Narva culture were also found to be more closely related with WHG, while the Pit–Comb Ware culture was more closely related to EHG. Northern and eastern areas of the eastern Baltic were found to be more closely related to EHG than southern areas. The study noted that EHGs, like SHGs and Baltic hunter-gatherers, carried high frequencies of the derived

alleles An allele (, ; ; modern formation from Greek ἄλλος ''állos'', "other") is a variation of the same sequence of nucleotides at the same place on a long DNA molecule, as described in leading textbooks on genetics and evolution. ::"The chrom ...

for SLC24A5 and SLC45A2, which are codings for

light skin Light skin is a human skin color that has a base level of eumelanin pigmentation that has adapted to environments of low UV radiation. Light skin is most commonly found amongst the native populations of Europe and East Asia as measured through ...

. Mathieson et al. (2018) analyzed the genetics of a large number of skeletons of prehistoric Eastern Europe. Thirty-seven samples were from Mesolithic and Neolithic Ukraine (9500-6000 BC). These were classified as intermediate between EHG and SHG. The males belonged exclusively to R haplotypes (particularly subclades of R1b1 and R1a) and I haplotypes (particularly subclades of I2). Mitochondrial DNA belonged almost exclusively to U (particularly subclades of U5 and U4). Dnieper–Donets culture

A large number of individuals from the Zvejnieki burial ground, which mostly belonged to the Kunda culture and Narva culture in the eastern Baltic, were analyzed. These individuals were mostly of WHG descent in the earlier phases, but over time EHG ancestry became predominant. The Y-DNA of this site belonged almost exclusively to haplotypes of haplogroup R1b1a1a and I2a1. The mtDNA belonged exclusively to haplogroup U (particularly subclades of U2, U4 and U5). Forty individuals from three sites of the

Iron Gates Mesolithic The Iron Gates Mesolithic is a Mesolithic archaeological culture, dating to between 13,000 and 6,000 years cal BCE, in the Iron Gates region of the Danube River, in modern Romania and Serbia. The people who inhabited the Iron Gates area during ...

in the

Balkans The Balkans ( ), also known as the Balkan Peninsula, is a geographical area in southeastern Europe with various geographical and historical definitions. The region takes its name from the Balkan Mountains that stretch throughout the who ...

were estimated to be of 85% WHG and 15% EHG descent. The males at these sites carried exclusively R1b1a and I (mostly subclades of I2a) haplotypes.

mtDNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial DNA ...

belonged mostly to U (particularly subclades of U5 and U4). People of the

Cucuteni–Trypillia culture The Cucuteni–Trypillia culture, also known as the Tripolye culture, is a Neolithic–Chalcolithic archaeological culture ( 5500 to 2750 BCE) of Eastern Europe. It extended from the Carpathian Mountains to the Dniester and Dnieper regions, cent ...

were found to harbor about 20% hunter-gatherer ancestry, which was intermediate between EHG and WHG. Narasimshan et al. (2019) coined a new ancestral component, West Siberian Hunter-Gatherer (WSHG). WSHGs contained about 30% EHG ancestry, 50% ANE ancestry, and 20% East Asian ancestry. Unlike the

, in the Dnieper–Donets culture no Caucasian Hunter-Gatherer (CHG) or Early European Farmer (EEF) ancestry has been detected. Dnieper-Donets males and Yamnaya males carry the same paternal haplogroups (R1b and I2a), suggesting that the CHG and EEF admixture among the Yamnaya came through EHG males mixing with EEF and CHG females. According to David W. Anthony, this suggests that the

were initially spoken by EHGs living in Eastern Europe.

Physical appearance

The EHGs are suggested to have had mostly

brown eyes Eye color is a polygenic phenotypic character determined by two distinct factors: the pigmentation of the eye's iris and the frequency-dependence of the scattering of light by the turbid medium in the stroma of the iris. In humans, the pig ...

and light skin, with "intermediate frequencies of the blue-eye variants" and "high frequencies of the light-skin variants." An EHG from Karelia was determined by to have high probabilities of being brown-eyed and dark haired, with a predicted light brown skin-pigmentation phenotype. Another EHG from Samara was predicted to be light skinned, and was determined to have a high probability of being green-eyed with a light hair shade, with a 75% calculated probability of being red. The rs12821256

allele An allele (, ; ; modern formation from Greek ἄλλος ''állos'', "other") is a variation of the same sequence of nucleotides at the same place on a long DNA molecule, as described in leading textbooks on genetics and evolution. ::"The chrom ...

of the

KITLG Stem cell factor (also known as SCF, KIT-ligand, KL, or steel factor) is a cytokine that binds to the c-KIT receptor (CD117). SCF can exist both as a transmembrane protein and a soluble protein. This cytokine plays an important role in hematopoie ...

gene that controls

melanocyte Melanocytes are melanin-producing neural crest-derived cells located in the bottom layer (the stratum basale) of the skin's epidermis, the middle layer of the eye (the uvea), the inner ear, vaginal epithelium, meninges, bones, and hear ...

development and

melanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...

synthesis, which is associated with

blond Blond (male) or blonde (female), also referred to as fair hair, is a hair color characterized by low levels of the dark pigment eumelanin. The resultant visible hue depends on various factors, but always has some yellowish color. The color ...

hair and first found in an individual from Siberia dated to around 15,000 BC, is found in three Eastern Hunter-Gatherers from Samara, Motala and Ukraine.

References

Bibliography

* * * * * * * * * * * * *

Research

Physical appearance

See also

References

Bibliography

Further reading