1. Introduction
Before DNA sequencing was developed around the turn of this century (you can see Wikipedia for a summary of the details: see especially "shotgun sequencing"), prehistory was a relatively blank slate. Scientists knew that anatomically modern humans had spread out of Africa, reaching Australia and the Americas after Eurasia. Carbon-14 dating in the 1950s let them start putting dates on the available fossils. And that was pretty much it. People could spread all sorts of weird theories, whether in academia or less reputably, and except for the most extraordinary claims, like huge lost civilizations in places that weren't flooded at the end of the last Ice Age, who could say?
Then it became possible to sequence a living or dead person's mitochondria, which is passed without recombination from mothers to children of both sexes, a male's Y-chromosome, passed without recombination from father to sons, even whole genomes. Suddenly evidence for who was doing what in prehistory got stronger.
2. Y-Haplogroups
Male lines A (Khoisan and West African forager) and B (Central African inc. Pygmies) are earliest African.
C-M130 is found at highest frequencies among indigenous people of Kazakhstan, Mongolia, Russian Far East, Polynesia, Australia.
D is found at high frequencies only in Andaman Islands, Tibet, Hokkaido, Sakhalin, Kamchatka. It was dominant among the high-density Jomon foragers of Japan. (Ainu = 1/3 Jomon-1/3 Okhotsk-1/3 rice farmer).
E1b1b1 appeared in Levant, was spread by Near East Neolithic colonists: North Africa, Ethiopia and Somalia 60-80%, Egypt 40%, Semitic-speakers (outside Africa), Cyprus, Sicily-South Italy and the Balkans 18-27%, except Kosovo 45%. Other E clades evolved in North Africa, E1b1a expanding with Niger river agriculture and later Bantu.
F split from GHIJK in India.
G appeared to the west and was spread by Neolithic colonists: up to 91% of 5000-3000 BC males from Central Germany, Alps, France, Catalonia. Now 5% there except Tyrolian Alps (40%), 5-13% Iran, 31% Georgia and up to 74% in Causasian minorities.
H exists at 25-40% in South India and Roma, otherwise spread at 2-10% from Levant by Neolithic colonists (5500 BC Linear Pottery Culture, Spain) and much later Khmer.
Basal IJ appeared in Iran (only place found today) >45,000 years ago.
I appeared in Europe by 31,300 years ago, associated with male survivors of the Last Glacial Maximum, became the majority lineage of European foragers, now highest in Dargwa (NE Caucasian) 58%, Croats up to 73% (Herzegovina, Hvar island), south Sweden 55-60%, Norway 45%.
J appeared between Yemen and Levant 43,000 years ago. J-M304* is rarely found outside island of Socotra (Yemen). Clade J2 later spread by Neolithic colonists, presently Ingush (NE Caucasian) 89%, Crete up to 44%, down to 2-10% in Scandinavia and North European plain, Central Asia, Niger river, Indian-colonized SE Asia. Clade J1 is strongly associated with Semitic-speakers at decreasing frequency from Yemen (76%), e.g. Iraqis 33%, African Arabs around 20%: pastoralist origin?
K split in India, K1/LT diversifying in place/spreading west, K2a (parent to NO) appearing somewhere between SE Asia and Omsk >45,000 years ago and K2b splitting into K2b1/MS (Melanesians/non-Andaman Negritos and 1/3 Australians) & P (ancestor of Native American Q and Eurasian R).
L existed by 4200 BC, found at 33% in Chalcolithic south Armenia (3% in Bronze Age Bactria). It has highest concentration in Karnataka (68% in one tribe), Tamil Nadu and spread almost nowhere east of India: 25-28% in Balochistan, northern Pashtuns, Kalasha, 55% in Parsi priests, drops to 1-5% in Danubian Europe, east France, England, Ireland, Arabian peninsula, Egypt. T is found in isolated pockets that peak in SE India and north Somalia.
N is North Eurasian: Samoyedic Uralic speakers up to 75-99%, Estonians and Saami 40%, Siberian Turkic peoples 27-84%, Tungusic-speakers up to 45%, Buryats 20-48%. Spread after Beringia disappeared. Uralic speakers arrived in the Baltic 500 BC.
O appeared between Bengal and the Mekong 34-39,000 years ago, later expanded with rice farmers. Note that most Austronesian ethnicities are dominant O but Polynesian founders merged with C-M38 east of Indonesia.
Q is widely distributed in Asia at rates of 2-10%, but 94% of Kets (Yenisei river near Altai), 66% of Selkups (Samoyedic), 60% of Chelkans (Turkic, Altai region). Oldest sample is >15,000 BC from Yenisei. 6 subclades founded the Americas: up to 94% of natives except in Alaska and much of Canada (80% Inuit and Na-Dene speakers besides Navajo). On the Asian side of the Bering Sea, 33% of Chukchi people, 39% of Yupik Eskimos.
R is known to have existed in Siberia by the Last Glacial Maximum, but both R1a and R1b show young expansions. Most R1a men are R1a1a1, which itself split in two estimated 5,800 years ago, with Europeans and Indo-Iranians having different clades (yet Asian clade Z93 has been found from 6,000 years ago in Ukraine). From 2900 BC, 75% of Corded Ware Culture men had European R1a1a1, while Asian diversification correlates with Indus Valley population boom around 2600 BC: now highest in Poles and some Sindhi/Punjabi/Gujarati groups (60-72%). R1b1a is first found in Venetia 14,100 years ago and now strongly concentrated in Western Europe (where Iberia has unique R1b3), but present in 100% of Yamnaya steppe samples (3300-2600 BC) and a Kura-Araxes sample (4000-2600 BC Caucasian).
Takeaways, working backwards from R to A:
The presence of R1a in 4000 BC Ukraine, highly diversified in Bronze-Age-to-now India, and none recovered from the burial mounds of Yamnaya or presumed successor cultures (Andronovo, etc.) partially demolishes the belief of most Indo-Europeanists that R1 distribution supports their pre-existing belief that IE and Hinduism were introduced by an invasive new element not a year earlier than 1500 BC. Hindu fundamentalists would prefer that it originated in India, but it looks more like a Western at-least-males element entered India around the time the wheel was introduced, which predates the Mature Indus Valley Civilization by about 700 years (~3300 BC in India as well as places as far off as Central Europe). In turn, this makes it look like Yamnaya and Andronovo were spreading a Central to Western European male lineage across the steppe after 3300 BC, though not to such an extreme as to prevent present-day Kazakhstan being deeply diverse.
R being sibling clade to Q (Native Americans) is not shocking, but non-obvious and pretty cool. The closest clade to that pair peaking (60%) in the short, woolly-haired Aeta people was a big surprise. I think it was once mainstream anthropological belief that Asian peoples the Spanish called Negritos were survivors of a southern Recent African Origin route and Caucasoids+other Asians+Native Americans had diversified out of the founders of the northern route.
The correlation between Y-haplogroup O and the spread of rice farming looks surprisingly simple. The genetics of cultures out-competed foragers by adopting the Fertile Crescent package is much more complicated.
I would have thought Finns (51-61% N), Saami and Estonians (40%) spread from the NE of Europe proper rather than their male lineage being close to people from tropical Asia.
The fact that L is associated by most researchers with Neolithic farmers and its distribution looks like this feels like fodder for some radical new Indo-European theory where it’s associated with the Neolithic Revolution (like Colin Renfrew) but came from India and was spread more by priests than wholesale demographic change. Its sibling clade T is even weirder.
The deep Paleolithic splits of I (Western forager) and J (Near East) from Indian K isn’t surprising, but it’s kind of funny that Paleolithic Europeans had their male lineage largely replaced by a mix of its sibling clade J, the older G, and descendants of an even older split close to most Black Africans[1]. And all this before another invasion brought the European demography of known history.
And the E1b1 Africans look like descendants of Neolithic colonists (at least male ones), modulo the continued presence of A and B. The suggestion that E never left Africa until E1b1b did so just in time to explain Natufian farmer DNA while C and D men jumped out of Africa so fast that it’s hard to find traces in Africa rather than places like Tibet and Hokkaido (D) seems like discarding Occam’s Razor to preserve a subtle ideology in the Out of Africa model.
3. Mitochondrial Haplogroups
The earliest split from Mitochrondrial Eve is L0 and L1-6. L0 is associated with the peopling of Southern Africa with ancestors of Khoisan groups. L1-6 is estimated (by Soares et al. 2009) 167±36 thousand years ago. Undifferentiated L1'2'3'4'5'6 has been found in Neanderthal fossils from the Caucasus (Mezmaiskaya cave) and Altai mountains (Denisova cave), dated to before 50,000 years ago. What's interesting about that is that L1 is estimated to have split off 140,000 years ago, and is most associated in living people with African Pygmies. This implies that Homo sapiens sapiens emigrated from Africa >140,000 years ago and interbred with Neanderthals without replacing them like in the more recent Out Of Africa event. L5 is another group mostly only found among Pygmies.
L2 diverged later than L5, perhaps 60-70,000 years ago (great job naming your groups, guys). Silva et al 2015 examined how it documents "60,000 years of interactions between Central and Eastern Africa". This haplogroup represents 70% of African maternal diversity, with the major exceptions being in the Sahara, Pygmies of the central rain forest, and Khoisan peoples. The latter two were displaced by the Bantu Expansion over the last few thousand years.
L6 is a rare haplogroup that reaches a peak of nearly 50% in Yemen and has its next-highest concentrations in the Horn of Africa. I cannot find research establishing how much this distribution might be due to the Paleolithic southern route Out of Africa vs. the archaeologically-documented interactions between Yemen and Ethiopia after 1000 BC, or continuous gene flow in between. An origin in the Arab slave trade seems highly unlikely, because the peak is in Yemen rather than Africa and so many African slaves were Bantu people.
L4 is concentrated in Tanzania among the Hadza at 60-83% and Sandawe at 48% (both peoples who were until recently foragers, Sandawe adopting livestock and farming earlier), declining in a gradient through east-central Africa.
L3 was carried by the last common ancestor (LCA) of all non-African humans, with a defining mutation ~70,000 years ago. It is unknown whether it originated in Asia or Africa: it has seven equidistant descendants, with the two clades of modern non-Africans being M and N.
Haplogroup N is older, sister clade M having a younger most recent common ancestor date than some subclades of N, such as haplogroup R. It has been proposed that N left Africa via the Northern route through the Levant, and M left Africa via Horn of Africa. "This theory was suggested because haplogroup N is by far the predominant haplogroup in Western Eurasia, and haplogroup M is absent in Western Eurasia, but is predominant in India and is common in regions East of India." However, "Southeast Asian populations and Indigenous Australians all possess deep rooted clades of both haplogroups M and N." (MacAulay et al 2005)
Haplogroup R* and R0 are found most commonly among the Soqotri, a Semitic-speaking people native to the island of Socotra, followed by Northeast Africa and the Arabian peninsula respectively. R0 has a daughter clade HV, which divides into V (Sami 40%), HV1 (Middle East), HV2 (South Asia), HV3 (Eastern Europe), and H (the most common mtDNA haplogroup in Europe).
R1 includes basal R1a* (found in Northwest Caucasian speakers and Brahmins from Uttar Pradesh), R1a1 (found in Northwest Caucasian speakers, Russians and Poles), R1b1 (Bulgarians, Armenians, Indians, Finland, and two Turkic peoples: Uyghurs and Yakuts).
R2 is found mainly in Pakistan's Balochistan province and NW India. It has a sister clade ancestral to J and T. T1a1a1 is particularly common in countries with high levels of Y-haplogroup R1a. In the Yamna culture, frequency of T1a and T2 were each 14.5%, a percentage higher than in any country today. T*, T1 and T2 are also found in Africa, primarily among Afro-Asiatic-speaking populations.
R5 is widely spread in India, peaking at 17%, with R5a1 and R5a2 found among Indians speaking Indo-Aryan and Dravidian languages respectively.
Haplogroup F is a daughter of R9. It is most common in East and Southeast Asia and has not been found among Native Americans. In India, it reaches 50% on the Nicobar islands and 31% in Arunachal Pradesh, the country's most Northeastern state, but is otherwise unknown.
Haplogroup B, daughter of R11, diversified in China, with Tianyuan man from a cave near Beijing, 42-39,000 years ago, having it. It is now most commonly seen in Southeast Asia and Austronesian speakers (indigenous Taiwanese, Polynesians, Madagascar, etc). Its subclade B4b is one of five haplogroups found among Native Americans, the others being A, C, D, and X. B, like X, is not found in Paleo-Siberian tribes of northeastern Siberia. It is however found elsewhere in Siberia: among Mongols, Tungusic-speakers, and Siberian Turkic peoples. It has also been found in Iraq.
Haplogroup P descended from R more than 50,000 years ago, when Papua New Guinea and Australia were first populated. It is also found among the Aeta (Philippine Negritos) at 40% and Melanesians, Malaysians, Indonesians.
Haplogroup U descended from R no later than ~45,000 years ago, as it was found in Paleolithic Western Siberian remains from that time. It is found in 15% of Indians (only 8% in tribal groups). It is found in approximately 11% of native Europeans and is held as the oldest maternal haplogroup found in that region. All but one ancient modern human sequences from Europe belonged to maternal haplogroup U, making it the dominant type of mtDNA in Europe before the spread of agriculture from Anatolia (Fu Qiaomei et al 2013). Africa has a unique subclade, U6, which peaks among Algerian Berbers (29%) and Egyptian Copts (27.6%). Prehistoric samples of U6 are common from the Iberomaurusian culture (final Paleolithic Morocco to Tunisia).
Haplogroup A has a low diversity of mitochondrial genomes that belong to it relative to the degree of divergence from its nearest outgroups in haplogroup N, suggesting a population bottleneck 20,000 years ago or less. It is found in 7-15% of Tibetans, Mongols, Koreans, Japanese and the majority of Inuit and many Native groups of North and Central America.
Haplogroup X arose from N >20,000 years ago and has a dramatically disjoint distribution. It is present among Native Americans (but not in South America) and remains from Morocco, Iberia, and Central Europe (Linear Pottery culture) circa 5,000 years ago, at rates similar to its modern 7% of Europeans. It reaches 27% among Druze. In North America it peaks among Algonquian (25%) and Sioux (15%).
Haplogroup N1a is believed to have mutated from N anywhere from 32,000 to 12,000 years ago, in the Near East and more specifically the northern Arabian peninsula. The younger date is close to the Pre-Pottery Neolithic of the Levant, an archaeological culture associated with the founder crops and domestic animals of western Eurasian agriculture and pastoralism. "The Linear Pottery Culture is credited with the first farming communities in Central Europe, marking the beginning of Neolithic Europe in the region some 7500 years ago. As of 2010, mitochondrial DNA analysis has been conducted on 42 specimens from five locations. Seven of these ancient individuals were found to belong to haplogroup N1a. A separate study analyzed 22 skeletons from European hunter-gatherer sites dated 13,400-2300 BC. Most of these fossils carried the mtDNA haplogroup U, which was not found in any of the Linear Pottery Culture sites. Conversely, N1a was not identified in any of the hunter-gatherer fossils, indicating a genetic distinction between Early European Farmers and late European hunter-gatherers. While no modern population is a close match to the LBK findings, the authors claim that the Linear Pottery population is most closely affiliated with modern Near East populations."
Haplogroup M (time depth >50,000 years) has the highest frequencies in India, Bangladesh, Nepal, China, Korea, and Japan, where frequencies range from 60%-80%. India has several M lineages that emerged directly from M*. Some small groups among eastern Siberians and Native Americans also have it at very high rates. Africa has only M1, raising questions of whether it appeared there or migrants from India eventually brought it west across Asia to Africa.
Haplogroup Q is a subclade of M29 with an inferred origin 50,000 years ago. It is especially found in New Guinea, Melanesia and Australian Aborigines. It is otherwise found, at much lower frequencies, among Austronesian-speaking peoples outside of Taiwan, indicating that both demic expansion and native foragers were important to the formation of the expanding Austronesians.
Haplogroup C is believed to have coalesced no later than 15,500 years ago, somewhere in Russia. It has a peak of 19-77% among different groups of the Tungusic-speaking Evenk people, with many Siberian Turkic peoples also having high frequencies. It drops to 15% or less among Mongols and Native Americans. In 2014, a now-extinct subclade was found in hunter-gatherers from Northwest Russia 7,500 years ago. C has a sister clade Z: Z1 is found at frequencies up to 28% among some Tungusic-speaking people and the Sami, while other subclades have found at 1-in-388 or less among Turks, Kazakhs, and Japanese.
Haplogroup D is another clade shared between Northeast Asia and the Americas. D4 is the most common maternal haplogroup for Japanese, Koreans, some Tungusic-speaking peoples, many Mongolic-speakers such as Buryats, and the Turkic Kazakhs and Telenghits. Up to 30% of Han Chinese have it. In the Americas, it predominates among fossils but rarely more than 17% among modern Natives except Paiute and Shoshone (up to 40%).
Haplogroup E is characteristic of Austronesian-speaking peoples, including native Taiwanese. The oldest evidence for it is from an 8,000-year-old skeleton found on Liang Island off the Southeast Chinese coast.
4. Prehistoric whole DNA
Beyond haplogroups, I've also written a summary of prehistoric whole-DNA sequencing from Europe. Other parts of the world to be added later.
4.1. Europe
As of 2014, some of the oldest European H. sapiens sapiens DNA was from Kostenki 14 in European Russia, 38,700 to 36,200 years ago (Seguin-Orlando et al).
“Kostenki 14 shares a close ancestry with the 24,000-year-old Mal’ta boy from central Siberia, European Mesolithic hunter-gatherers, some contemporary western Siberians, and many Europeans, but not eastern Asians. Additionally, the Kostenki 14 genome shows evidence of shared ancestry with a population basal to all Eurasians that also relates to later European Neolithic farmers. We find that Kostenki 14 contains more Neandertal DNA that is contained in longer tracts than present Europeans.”
Before 38,700 years ago, you’ll see claims that all non-Africans were one undifferentiated population. I suspect that may be incorrect, there being evidence of Saharan and boat routes Out Of Africa.
European early modern humans (EEMH) lineages of 39 -26,000 years ago (often called Aurignacian after an archaeological horizon) were still part of a large Western Eurasian “meta-population”, related to Central and Western Asian populations. I would surmise that these people should be reconstructed as Cro-Magnon skeletons (as they used to be called: now EEMH) with skin like South Indians of farmer descent (i.e. native Dravidian speakers), except for a northern clinal variation toward fair skin and the red hair of European Neanderthals (not to be confused with West Asian Neanderthals, from whom I don’t think we have any red hair markers).
About that skin depigmentation:
“Using compound haplotype systems consisting of rapidly evolving microsatellites linked to one single-nucleotide polymorphism in each gene, we estimate that the onset of the sweep shared by Europeans and East Asians at KITLG occurred approximately 30,000 years ago, after the out-of-Africa migration, whereas the selective sweeps for the European-specific alleles at TYRP1, SLC24A5, and SLC45A2 started much later, within the last 11,000–19,000 years, well after the first migrations of modern humans into Europe.” (Beleza et al 2012) I don’t uncritically agree with this, because that would imply all EEMH redheads, who had “more Neandertal DNA that is contained in longer tracts than present”, being as dark-skinned as tropical East Asians or Dravidian-speaking peoples, lacking all of TYRP1, SLC24A5, and SLC45A2 in the time between our ~39,000 year-old sample and 19,000 years ago or later.
Anyway, as de-glaciation commenced in the Northern Hemisphere ~19,000 years ago, we find founder effects producing the lineage dubbed West European Hunter-Gatherer, which emerged from the Solutrean refugium of the Last Glacial Maximum (Jones et al 2015).
“We find that Caucasus hunter-gatherers (CHG) belong to a distinct ancient clade that split from western hunter-gatherers ∼45,000 years ago, shortly after the expansion of anatomically modern humans into Europe and from the ancestors of Neolithic farmers ∼25,000 years ago, around the Last Glacial Maximum. CHG genomes significantly contributed to the Yamnaya steppe herders…” (about whom more later).
All of the successfully tested Mesolithic WHG Y-chromosomes, one from Luxembourg and four from Motala, Sweden, belonged to haplogroup I. Haplogroup I is the main candidate for Europe’s indigenous Y-haplogroup, which is today the most common Y-haplogroup in most of Scandinavia.
The DNA that’s been extracted from prehistoric farmer skeletons indicates that the Neolithic Revolution in Europe was mostly a story of population replacement, not adoption of new technology. Non-Nordic lineages of WHG appear to have made minimal contributions to the descendants of the invading farmers, who cluster with modern Aegean people: but bear in mind that historical Greeks have a para-Nordic WHG element (Homer mentions white-skinned, red-haired Mycenaean aristoi). For this period, think Native American ancestry as a percentage of heritage among farmers north of the Rio Grande in recent times.
That Nordic WHG heritage survived among Europeans seems to be thanks to the Fertile Crescent agricultural package hitting a wall as it approached the Baltic. The “Danubian cultures” archaeological group produce Early European Farmer bones, and as you can see in yellow on this map, they didn’t reach the Baltic and touch the North Sea only at the mouth of the Rhine. Erteboelle and Comb Ceramic Culture on the same map represent hunter-gatherers selectively adopting technology from the invaders: if they didn’t already have villages, they were producing pottery and settling down, but they were more into intensive fishing than agriculture. It seems that domesticated species of the Fertile Crescent package needed time to be bred for colder conditions, giving the Nordic WHGs time to survive the Neolithic Revolution.
At ~8,400 years ago, the Early European Farmers (EEF), very closely related the Anatolian or Levantine farmers (other offshoots of which are colonizing Mesopotamia and the Nile/Green Sahara), are colonizing the northern margin of Greece*, places like Epirus and Corfu. 1-4 centuries later, their “Cardium pottery culture” is in modern Albania, Croatia, and the Adriatic coast of Italy. Contemporary with this, early examples of their pottery appear in Sardinia, though I don’t know when their “race” colonized the island (21st century Sardinians are >50% EEF). At 5500 cal. BC, the Cardium pottery culture expands into the southern half of France and parts of Spain.
*They were in the rest of mainland Greece and Crete somewhat earlier.
Analysis for ancient DNA found the rare mtDNA basal haplogroup N*, supporting an early Neolithic maritime colonization of Mainland Europe through Cyprus and the Aegean Islands by Near-Eastern farmers (Fernandez et al 2014).
By the time farmers reached Spain, they would have encountered a branch of WHGs with darker skin than themselves and the recessive gene for blue eyes. Maybe before that: I need data from outside Iberia.
Meanwhile, the Danubian or Linear Pottery Culture (which you’ll encounter abbreviated as LBK) spread from the Hungarian Danube (before 5600 BC) to Austria, central Germany and central Poland circa 5,500 BC, spreading east and north over the next three centuries until they hit the aforementioned wall for their domesticates very near the Baltic and somewhat further from the North Sea except along the Rhine. At the former limit is where they would have interacted with Nordic WHGs without replacing them.
To conclude this summary of the European Neolithic west of the Black Sea, also at circa 5300-5200 BC we find multiple cultures interacting in the Netherlands. Probably the most significant is the Ertebølle-Ellerbek horizon, which had an all-but identical southern form in Limburg, the Netherlands in contact with LBK. Ertebølle-Ellerbek skeletal remains are meager, but some DNA sequences have been collected, showing genetic links between Limburg, northern Germany, Denmark, and peninsular Scandinavia. Farmers make the jump from the mouth of the Rhine or northern France to the British Isles circa 4000 BC, and this population was >50% Early European Farmer (This raises the question of whether they were also <50% para-Nordic WHG like the Luxembourg Mesolithic sample. I still need to find Mesolithic Britain/Ireland/north France data.)
Looking east of the Black Sea, unter-gatherers of the Caucasus split off from the European hunter-gatherers who would go on to become dark-skinned, blue-eyed Iberians, light-skinned, redheaded Nordics, etc ~45,000 years ago. Until the Last Glacial Maximum ~25,000 years ago, they remained part of the Anatolian HG population whose descendants would include the first wheat farmers. Post-isolation, they are dubbed CHG, for Caucasian Hunter-Gatherer. Jones et al. 2015 analyzed CHG ancestry as represented by a Upper Palaeolithic male from Satsurblia cave, and a Mesolithic one from Kotias Klde cave, both in the foothills of western Georgia. These two males carried Y-DNA haplogroup J* and J2a. The researchers found that these Caucasus hunters were probably the source of the Near Eastern DNA in the Yamnaya (who I’ll finally get to anon).
Let’s emphasize “probably” there: this is one paper demurring to claim very high Bayesian probability. A competing claim is that Near Eastern DNA made its way to the Eurasian steppe via farmers from Iran.
Interesting to note that DNA from the Maykop farming culture of the north Caucasus has been sequenced and found to not be related to the adjacent steppe population.
Something amazing happened around the middle of the 4th millennium BC. Pictographs of wheeled vehicles appear on clay tablets from Uruk in modern south Iraq dated between 3700-3500 BC. Between 3500-3350 BC, evidence of the wheel suddenly appears all the way from Harappa in India (Ravi phase) to southern Poland (Funnelbeaker culture). We have every reason to suspect that it was invented by a speaker of proto-Indo-European or proto-Semitic, as the common ancestor of all European, Iranian and Indian wheel words is reconstructed by experts as *kʷékʷlos, a proper grammatical “reduplicated derivative” of *kʷel- (“to turn”). It’s also reconstructed as galgal, simply “roll” reduplicated, in Semitic languages. In Sumerian, “chariot” was GIGIR, with no known native source. “Wheel” is *grgar in the reconstructed ancestor of Georgian and the other South Caucasian languages, which likewise looks like a loan.
Unfortunately, the epistemology of dating in archaeology and genetics are not identical, so we don’t know if the people who first introduced the wheel to India and Eastern Europe changed the areas’s genetic profiles. But change did come from pastoraists who knew how to build wheeled vehicles.
As we saw, the most common Y-chromosomal haplogroup in Europe is R1b, with the closely-related R1a a contender for third place after the believed-indigenous (i.e. WHG had it) I1. Here’s a map. As the Bronze Age was starting in civilization to the south, one of the early adopters in the illiterate north was a nomadic culture called Yamnaya. They had carts, to which they yoked horses, a domesticate then unknown in Mesopotamia. They buried elite males with weapons under artificial hills, which we call kurgans. And when DNA from their skeletons is sequenced, the most common Y-haplogroup is R1b. Scientific consensus is that the spread of these genes was caused by the same phenomenon documented archaeologically as the collapse of prehistoric (i.e. illiterate) farming cultures and replacement by pastoralist cultures in Eastern Europe south of the Baltic and north of the Greek peninsula.
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