Proto-Anatolian: The Precursor of Proto-Indo-European (PIE)

If Indo-Anatolian was indeed influenced by a North Caucasian substrate, it
stands to reason that its ancestor moved into the steppes north of the
Caucasus from somewhere else. The Uralic connection suggests that this
somewhere else must be sought more to the north and/or the east. As for the
question when Proto-Indo-Uralic was spoken, only a very rough estimate is
possible on the basis of the rather limited number of etymological
correspondences between the basic lexicons of Indo-European and Uralic.
The relatively low number of probable cognates suggests that at least a couple
of millennia must have passed between the dissolution of Proto-Indo-Uralic
and its daughter languages Proto-Indo-Anatolian and Proto-Uralic (Napol’skix
1997: 143), even if we take into account the temporal gap of up to 2500 years
between Proto-Indo-Anatolian and Proto-Uralic. Proto-Indo-Uralic would
then have been spoken in or around the 7th millennium BCE.



We can conclude that ‘core’ Proto-Indo-European, consisting of the
Brugmannian branches of Indo-European, had a number of precursors. Its
direct ancestor was Proto-Indo-Tocharian, to which it was very similar. A
more distant, but still clearly recognizable ancestor was Proto-Indo-Anatolian,
which seems to have been spoken at least a thousand years earlier. Finally,
there was a distant Proto-Indo-Uralic ancestor, with which ‘core’
Proto-Indo-European shared only a limited number of words and a few dozen
grammatical characteristics and which must have been spoken at least several
millennia before Proto-Indo-Anatolian. The papers in this volume reflect the
state of the art in the research into these ancestors of ‘core’
Proto-Indo-European.


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The Anatolian Neolithic is a likely source of the European Neolithic
Our population sample from the Neolithic of Northwestern Anatolia (Fig. 1, Extended Data
Fig. 2) has a clear affinity to early European farmers. However, EEF are shifted slightly
towards the direction of the WHG in the PCA (Fig. 1) and share more alleles with them than
do the Anatolian Neolithic samples (Extended Data Table 3).
“World Foci 15” set of outgroups: Ami, Biaka, Bougainville, Chukchi, Eskimo, Han, Ju_hoan_North,
Karitiana, Kharia, Mbuti, Onge, Papuan, She, Ulchi, Yoruba
To enhance this set’s ability to differentiate between reference populations, we can add to it
populations that are differentially related to them. For example, when we study admixture in
LBK_EN between Anatolia_Neolithic and WHG, we can add EHG as an additional outgroup,
as f4(WHG, Anatolia_Neolithic; EHG, Mbuti)= 0.00823 (Z=22.2), so allele sharing with EHG
helps distinguish between WHG and Anatolia_Neolithic as sources of ancestry.
Extended Data Fig. 3C shows that Early Neolithic groups from Europe can be modeled as
predominantly of Anatolia_Neolithic ancestry with only 7-11% WHG ancestry. This confirms
the visual impression from the PCA (Fig. 1) and ADMIXTURE analysis (Extended Data Fig.
2) of the close relationship between Early Neolithic Europe and Neolithic Anatolia. The
relationship is also supported by the low FST=0.005±0.00046 (Supplementary Data Table 2)
between LBK_EN and Anatolia_Neolithic and 0.006±0.00054 between Hungary_EN and
Anatolia_Neolithic. A direct link between Neolithic Europe and Anatolia is furnished by the
fact that around half of the Y-chromosomes of Anatolia_Neolithic belong to haplogroup G2a
typical of ancient EEF samples5,20, and by the occurrence of mtDNA haplogroup N1a which is
typical of EEF21,22 in Neolithic Anatolia (Supplementary Data Table 1).
Our results add to the evidence in favor of the hypothesis that the Anatolian Neolithic was the
source of the European Neolithic, but we add two notes of caution. First, the fact that our
samples are from northwestern Anatolia should not be taken to imply that the Neolithic must
have entered Europe from that direction: it simply places a plausible ancestor population at
the doorstep of Europe and confirms the hypothesis that the genetic similarity between central
European and Iberian farmers could be explained by such a common ancestor. Second, we do
not know the geographical extent of people with this ancestry within Anatolia itself and
across the Near East. If the population of which the Anatolian Neolithic was a part was
sufficiently widespread this ancestry could have entered Europe from a different geographic
route. To understand the geographic spread and deeper origin of the Anatolian farmers, it will
be necessary to obtain ancient genomes from multiple locations within the Near East.

Lack of steppe ancestry in the Iberian Chalcolithic
Ancient North Eurasian ancestry is ubiquitous across present-day Europe5,7 but was absent in
Early and Middle Neolithic Europe5, raising the question of when it spread there. When we
add Yamnaya_Samara as a third ancestral population there is no improvement in fit for the
Iberia_Chalcolithic population (Extended Data Fig. 3B), and this population can be modeled
as a mixture of ~77%/23% Anatolia_Neolithic/WHG (Extended Data Fig. 3C). The Iberian
Chalcolithic did not have more WHG ancestry than the earlier Middle Neolithic population,
as the statistic f4(Iberia_MN, Iberia_Chalcolithic; WHG, Chimp) is not significantly different
from zero (Extended Data Table 3). A recent analysis of a different Iberian Chalcolithic
population also suggests that it had more hunter-gatherer ancestry than the earlier European
farmers4.
The Iberian Chalcolithic population lacks steppe ancestry, but Late Neolithic central and
northern Europeans have substantial such ancestry (Extended Data Fig. 3E) suggesting that
the spread of ANE/steppe ancestry did not occur simultaneously across Europe. All presentday Europeans have less steppe ancestry than the Corded Ware5, suggesting that this ancestry was diluted as the earliest descendants of the steppe migrants admixed with local populations.
However, the statistic f4(Basque, Iberia_Chalcolithic; Yamnaya_Samara,Chimp)=0.00168 is
significantly positive (Z=8.1), as is the statistic f4(Spanish, Iberia_Chalcolithic;
Yamnaya_Samara, Chimp)= 0.00092 (Z=4.6). This indicates that steppe ancestry occurs in
present-day southwestern European populations, and that even the Basques cannot be
considered as mixtures of early farmers and hunter-gatherers without it4.