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the lioness,
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http://biorxiv.org/content/early/2016/06/16/059311

The genetic structure of the world's first farmers

2016 Iosif Lazaridis, et al


We report genome-wide ancient DNA from 44 ancient Near Easterners ranging in time 86 between ~12,000-1,400 BCE, from Natufian hunter-gatherers to Bronze Age farmers. 87 We show that the earliest populations of the Near East derived around half their 88 ancestry from a ‘Basal Eurasian’ lineage that had little if any Neanderthal admixture 89 and that separated from other non-African lineages prior to their separation from each 90 other. The first farmers of the southern Levant [Israel and Jordan] and Zagros
91 Mountains [Iran] were strongly genetically differentiated, and each descended from 92 local hunter-gatherers. By the time of the Bronze Age, these two populations and 93 Anatolian-related farmers had mixed with each other and with the hunter-gatherers of 94 Europe to drastically reduce genetic differentiation. The impact of the Near Eastern 95 farmers extended beyond the Near East: farmers related to those of Anatolia spread 96 westward into Europe; farmers related to those of the Levant spread southward into 97 East Africa; farmers related to those from Iran spread northward into the Eurasian 98 steppe; and people related to both the early farmers of Iran and to the pastoralists of 99 the Eurasian steppe spread eastward into South Asia.
100 Between 10,000-9,000 BCE, humans began practicing agriculture in the Near East1. In the 101 ensuing five millennia, plants and animals domesticated in the Near East spread throughout 102 West Eurasia [a vast region that also includes Europe] and beyond. The relative homogeneity 103 of present-day West Eurasians in a world context2 suggests the possibility of extensive 104 migration and admixture that homogenized geographically and genetically disparate sources 105 of ancestry. The spread of the world’s first farmers from the Near East would have been a 106 mechanism for such homogenization. To date, however, due to the poor preservation of DNA 107 in warm climates, it has been impossible to study the population structure and history of the 108 first farmers and to trace their contribution to later populations.
109 In order to overcome the obstacle of poor DNA preservation, we took advantage of two 110 methodological developments. First, we sampled from the inner ear region of the petrous 111 bone3,4 that can yield up to ~100 times more endogenous DNA than other skeletal elements4. 112 Second, we used in-solution hybridization5 to enrich extracted DNA for about 1.2 million 113 single nucleotide polymorphism [SNP] targets6,7, making efficient sequencing practical by 114 filtering out microbial and non-informative human DNA. We merged all sequences extracted 115 from each individual, and randomly sampled a single sequence to represent each SNP, 116 restricting to individuals with at least 9,000 SNPs covered at least once. We obtained 117 genome-wide data passing quality control for 45 individuals on whom we had a median
3
bioRxiv preprint first posted online Jun. 16, 2016; doi: http://dx.doi.org/10.1101/059311. The copyright holder for this preprint [which was not peer-reviewed] is the author/funder. It is made available under a CC-BY-NC-ND 4.0 International license.
118 coverage of 172,819 SNPs [Methods]. We assembled radiocarbon dates for 26 individuals 119 [22 new generated for this study] [Supplementary Data Table 1].
120 The newly reported ancient individuals date to ~12,000-1,400 BCE and come from the 121 southern Caucasus [Armenia], northwestern Anatolia [Turkey], Iran, and the southern Levant 122 [Israel and Jordan] [Supplementary Data Table 1, Fig. 1a]. [One individual had a radiocarbon 123 date that was not in agreement with the date of its archaeological context and was also a 124 genetic outlier.] The samples include Epipaleolithic Natufian hunter-gatherers from Raqefet 125 Cave in the Levant [12,000-9,800 BCE]; a likely Mesolithic individual from Hotu Cave in the 126 Alborz mountains of Iran [probable date of 9,100-8,600 BCE]; Pre-Pottery Neolithic farmers 127 from ‘Ain Ghazal and Motza in the southern Levant [8,300-6,700 BCE]; and early farmers 128 from Ganj Dareh in the Zagros mountains of western Iran [8,200-7,600 BCE]. The samples 129 also include later Neolithic, Chalcolithic [~4,800-3,700 BCE], and Bronze Age [~3,350- 130 1,400 BCE] individuals [Supplementary Information, section 1]. We combined our data with 131 previously published ancient data7,8,9,10,8,10-15 to form a dataset of 281 ancient individuals. We 132 then further merged with 2,583 present-day people genotyped on the Affymetrix Human 133 Origins array13,16 [238 new] [Supplementary Data Table 2; Supplementary Information, 134 section 2]. We grouped the ancient individuals based on archaeological culture and 135 chronology [Fig. 1a; Supplementary Data Table 1]. We refined the grouping based on 136 patterns evident in Principal Components Analysis [PCA]17 [Fig. 1b; Extended Data Fig. 1], 137 ADMIXTURE model-based clustering18 [Fig. 1c], and ‘outgroup’ f3-analysis [Extended Data 138 Fig. 2]. We used f4-statistics to identify outlier individuals and to cluster phylogenetically 139 indistinguishable groups into ‘Analysis Labels’ [Supplementary Information, section 3].
140 We analyzed these data to address six questions. [1] Previous work has shown that the first 141 European farmers harboured ancestry from a Basal Eurasian lineage that diverged from the 142 ancestors of north Eurasian hunter-gatherers and East Asians before they separated from each 143 other13 What was the distribution of Basal Eurasian ancestry in the ancient Near East? [2]
144 Were the first farmers of the Near East part of a single homogeneous population, or were they 145 regionally differentiated? [3] Was there continuity between late pre-agricultural hunter- 146 gatherers and early farming populations, or were the hunter-gatherers largely displaced by a 147 single expansive population as in early Neolithic Europe?8 [4] What is the genetic
148 contribution of these early Near Eastern farmers to later populations of the Near East? [5] 149 What is the genetic contribution of the early Near Eastern farmers to later populations of
4
bioRxiv preprint first posted online Jun. 16, 2016; doi: http://dx.doi.org/10.1101/059311. The copyright holder for this preprint [which was not peer-reviewed] is the author/funder. It is made available under a CC-BY-NC-ND 4.0 International license.
150 mainland Europe, the Eurasian steppe, and to populations outside West Eurasia? [6] Do our 151 data provide broader insights about population transformations in West Eurasia?
152 Basal Eurasian ancestry was pervasive in the ancient Near East and associated with 153 reduced Neanderthal ancestry
154 The ‘Basal Eurasians’ are a lineage hypothesized13 to have split off prior to the differentiation 155 of all other Eurasian lineages, including both eastern non-African populations like the Han 156 Chinese, and even the early diverged lineage represented by the genome sequence of the 157 ~45,000 year old Upper Paleolithic Siberian from Ust’-Ishim11. To test for Basal Eurasian 158 ancestry, we computed the statistic f4[Test, Han; Ust’-Ishim, Chimp] [Supplementary
159 Information, section 4], which measures the excess of allele sharing of Ust’-Ishim with a 160 variety of Test populations compared to Han as a baseline. This statistic is significantly 161 negative [Z<-3.7] for all ancient Near Easterners as well as Neolithic and later Europeans, 162 consistent with their having ancestry from a deeply divergent Eurasian lineage that separated 163 from the ancestors of most Eurasians prior to the separation of Han and Ust’-Ishim. We used 164 qpAdm7 to estimate Basal Eurasian ancestry in each Test population. We obtain the highest 165 estimates in the earliest populations from both Iran [66±13% in the likely Mesolithic sample, 166 48±6% in Neolithic samples], and the Levant [44±8% in Epipaleolithic Natufians] [Fig. 2], 167 showing that Basal Eurasian ancestry was widespread across the ancient Near East.
168 West Eurasians harbour significantly less Neanderthal ancestry than East Asians19,20-23, which 169 could be explained if West Eurasians [but not East Asians] have partial ancestry from a 170 source diluting their Neandertal inheritance21. Supporting this theory, we observe a negative 171 correlation between Basal Eurasian ancestry and the rate of shared alleles with Neanderthals19 172 [Supplementary Information, section 5; Fig. 2]. By extrapolation, we infer that the Basal
173 Eurasian population had lower Neanderthal ancestry than non-Basal Eurasian populations and 174 possibly none [ninety-five percent confidence interval truncated at zero of 0-60%; Fig. 2; 175 Methods]. The finding of little if any Neanderthal ancestry in Basal Eurasians could be 176 explained if the Neanderthal admixture into modern humans 50,000-60,000 years ago11
177 largely occurred after the splitting of the Basal Eurasians from other non-Africans.
178 It is striking that the highest estimates of Basal Eurasian ancestry are from the Near East, 179 given the hypothesis that it was there that most admixture between Neanderthals and modern 180 humans occurred19,24. This could be explained if Basal Eurasians thoroughly admixed into the 181 Near East before the time of the samples we analyzed but after the Neanderthal admixture.
5
bioRxiv preprint first posted online Jun. 16, 2016; doi: http://dx.doi.org/10.1101/059311. The copyright holder for this preprint [which was not peer-reviewed] is the author/funder. It is made available under a CC-BY-NC-ND 4.0 International license.
182 Alternatively, the ancestors of Basal Eurasians may have always lived in the Near East, but 183 the lineage of which they were a part did not participate in the Neanderthal admixture.
184 A population without Neanderthal admixture, basal to other Eurasians, may have plausibly 185 lived in Africa. Craniometric analyses have suggested that the Natufians may have migrated 186 from north or sub-Saharan Africa25,26, a result that finds some support from Y chromosome 187 analysis which shows that the Natufians and successor Levantine Neolithic populations
188 carried haplogroup E, of likely ultimate African origin, which has not been detected in other 189 ancient males from West Eurasia [Supplementary Information, section 6] 7,8. However, no 190 affinity of Natufians to sub-Saharan Africans is evident in our genome-wide analysis, as 191 present-day sub-Saharan Africans do not share more alleles with Natufians than with other 192 ancient Eurasians [Extended Data Table 1]. [We could not test for a link to present-day North 193 Africans, who owe most of their ancestry to back-migration from Eurasia27,28.] The idea of 194 Natufians as a vector for the movement of Basal Eurasian ancestry into the Near East is also 195 not supported by our data, as the Basal Eurasian ancestry in the Natufians [44±8%] is
196 consistent with stemming from the same population as that in the Neolithic and Mesolithic 197 populations of Iran, and is not greater than in those populations [Supplementary Information, 198 section 4]. Further insight into the origins and legacy of the Natufians could come from 199 comparison to Natufians from additional sites, and to ancient DNA from north Africa.
200 Extreme regional differentiation in the ancient Near East
201 PCA on present-day West Eurasian populations [Methods] [Extended Data Fig. 1] on which 202 we projected the ancient individuals [Fig. 1b] replicates previous findings of a Europe-Near 203 East contrast along the horizontal Principal Component 1 [PC1] and parallel clines [PC2] in 204 both Europe and the Near East [Extended Data Fig. 1]7,8,13. Ancient samples from the Levant 205 project at one end of the Near Eastern cline, and ancient samples from Iran at the other. The 206 two Caucasus Hunter Gatherers [CHG]9 are less extreme along PC1 than the Mesolithic and 207 Neolithic individuals from Iran, while individuals from Chalcolithic Anatolia, Iran, and
208 Armenia, and Bronze Age Armenia occupy intermediate positions. Qualitatively, the PCA 209 has the appearance of a quadrangle whose four corners are some of the oldest samples: 210 bottom-left: Western Hunter Gatherers [WHG], top-left: Eastern Hunter Gatherers [EHG], 211 bottom-right: Neolithic Levant and Natufians, top-right: Neolithic Iran. This suggests the 212 hypothesis that diverse ancient West Eurasians can be modelled as mixtures of as few as four
6
bioRxiv preprint first posted online Jun. 16, 2016; doi: http://dx.doi.org/10.1101/059311. The copyright holder for this preprint [which was not peer-reviewed] is the author/funder. It is made available under a CC-BY-NC-ND 4.0 International license.
213 streams of ancestry related to these populations, which we confirmed using qpWave7 214 [Supplementary Information, section 7].
215 We computed squared allele frequency differentiation between all pairs of ancient West 216 Eurasians29 [Methods; Fig. 3; Extended Data Fig. 3], and found that the populations at the 217 four corners of the quadrangle had differentiation of FST=0.08-0.15, comparable to the value 218 of 0.09-0.13 seen between present-day West Eurasians and East Asians [Han] 219 [Supplementary Data Table 3]. In contrast, by the Bronze Age, genetic differentiation 220 between pairs of West Eurasian populations had reached its present-day low levels [Fig. 3]: 221 today, FST is ≤0.025 for 95% of the pairs of West Eurasian populations and ≤0.046 for all 222 pairs. These results point to a demographic process that established high differentiation 223 across West Eurasia and then reduced this differentiation over time.
224 Continuity between pre-farming hunter-gatherers and early farmers of the Near East
225 Our data document continuity across the hunter-gatherer / farming transition, separately in 226 the southern Levant and in the southern Caucasus-Iran highlands. The qualitative evidence 227 for this is that PCA, ADMIXTURE, and outgroup f3 analysis cluster Levantine hunter- 228 gatherers [Natufians] with Levantine farmers, and Iranian and Caucasus Hunter Gatherers 229 with Iranian farmers [Fig. 1b; Extended Data Fig. 1; Extended Data Fig. 2]. We confirm this 230 in the Levant by showing that its early farmers share significantly more alleles with Natufians 231 than with the early farmers of Iran: the statistic f4[Levant_N, Chimp; Natufian, Iran_N] is
232 significantly positive [Z=13.6]. The early farmers of the Caucasus-Iran highlands similarly 233 share significantly more alleles with the hunter-gatherers of this region than with the early 234 farmers from the Levant: the statistic f4[Iran_N, Chimp; Caucasus or Iran highland hunter- 235 gatherers, Levant_N] is significantly positive [Z>6].
236 How diverse first farmers of the Near East mixed to form the region’s later populations
237 Almost all ancient and present-day West Eurasians have evidence of significant admixture 238 between two or more ancestral populations, as documented by statistics of the form f3[Test; 239 Reference1, Reference2] which if negative, show that a Test population’s allele frequencies 240 tend to be intermediate between two Reference populations16 [Extended Data Table 2]. To 241 better understand the admixture history beyond these patterns, we used qpAdm7, which can 242 evaluate whether a particular Test population is consistent with being derived from a set of 243 proposed source populations, and if so, infer mixture proportions [Methods]. We used this
7
bioRxiv preprint first posted online Jun. 16, 2016; doi: http://dx.doi.org/10.1101/059311. The copyright holder for this preprint [which was not peer-reviewed] is the author/funder. It is made available under a CC-BY-NC-ND 4.0 International license.
244 approach to carry out a systematic survey of ancient West Eurasian populations to explore 245 their possible sources of admixture [Fig. 4; Supplementary Information, section 7].
246 Among first farmers, those of the Levant trace ~2/3 of their ancestry to people related to 247 Natufian hunter-gatherers and ~1/3 to people related to Anatolian farmers [Supplementary 248 Information, section 7]. Western Iranian first farmers cluster with the likely Mesolithic 249 HotuIIIb individual and more remotely with hunter-gatherers from the southern Caucasus 250 [Fig. 1b], and share alleles at an equal rate with Anatolian and Levantine early farmers 251 [Supplementary Information, section 7], highlighting the long-term isolation of western Iran.
252 During subsequent millennia, the early farmer populations of the Near East expanded in all 253 directions and mixed, as we can only model populations of the Chalcolithic and subsequent 254 Bronze Age as having ancestry from two or more sources. The Chalcolithic people of western 255 Iran can be modelled as a mixture of the Neolithic people of western Iran, the Levant, and 256 Caucasus Hunter Gatherers [CHG], consistent with their position in the PCA [Fig. 1b].
257 Admixture from populations related to the Chalcolithic people of western Iran had a wide 258 impact, consistent with contributing ~44% of the ancestry of Levantine Bronze Age 259 populations in the south and ~33% of the ancestry of the Chalcolithic northwest Anatolians in 260 the west. Our analysis show that the ancient populations of the Chalcolithic Iran, Chalcolithic 261 Armenia, Bronze Age Armenia and Chalcolithic Anatolia were all composed of the same 262 ancestral components, albeit in slightly different proportions [Fig. 4b; Supplementary 263 Information, section 7].
264 The Near Eastern contribution to Europeans, East Africans and South Asians
265 Admixture did not only occur within the Near East but extended towards Europe. To the 266 north, a population related to people of the Iran Chalcolithic contributed ~43% of the 267 ancestry of early Bronze Age populations of the steppe. The spread of Near Eastern ancestry 268 into the Eurasian steppe was previously inferred7 without access to ancient samples, by 269 hypothesizing a population related to present-day Armenians as a source7,8. To the west, the 270 early farmers of mainland Europe were descended from a population related to Neolithic 271 northwestern Anatolians8. This is consistent with an Anatolian origin of farming in Europe, 272 but does not reject other sources, since the spatial distribution of the Anatolian/European-like 273 farmer populations is unknown. We can rule out the hypothesis that European farmers stem 274 directly from a population related to the ancient farmers of the southern Levant30,31, however,
8
bioRxiv preprint first posted online Jun. 16, 2016; doi: http://dx.doi.org/10.1101/059311. The copyright holder for this preprint [which was not peer-reviewed] is the author/funder. It is made available under a CC-BY-NC-ND 4.0 International license.
275 since they share more allele with Anatolian Neolithic farmers than with Levantine farmers as 276 attested by the positive statistic f4[Europe_EN, Chimp; Anatolia_N, Levant_N] [Z=15].
277 Migrations from the Near East also occurred towards the southwest into East African 278 populations which experienced West Eurasian admixture ~1,000 BCE32,33. Previously, the 279 West Eurasian population known to be the best proxy for this ancestry was present-day 280 Sardinians33, who resemble Neolithic Europeans genetically13,34. However, our analysis 281 shows that East African ancestry is significantly better modelled by Levantine early farmers 282 than by Anatolian or early European farmers, implying that the spread of this ancestry to East 283 Africa was not from the same group that spread Near Eastern ancestry into Europe [Extended 284 Data Fig. 4; Supplementary Information, section 8].
285 In South Asia, our dataset provides insight into the sources of Ancestral North Indians [ANI], 286 a West Eurasian related population that no longer exists in unmixed form but contributes a 287 variable amount of the ancestry of South Asians35,36 [Supplementary Information, section 9] 288 [Extended Data Fig. 4]. We show that it is impossible to model the ANI as being derived
289 from any single ancient population in our dataset. However, it can be modelled as a mix of 290 ancestry related to both early farmers of western Iran and to people of the Bronze Age 291 Eurasian steppe; all sampled South Asian groups are inferred to have significant amounts of 292 both ancestral types. The demographic impact of steppe related populations on South Asia 293 was substantial, as the Mala, a south Indian population with minimal ANI along the ‘Indian 294 Cline’ of such ancestry35,36 is inferred to have ~18% steppe-related ancestry, while the Kalash 295 of Pakistan are inferred to have ~50%, similar to present-day northern Europeans7.
296 Broader insights into population transformations across West Eurasia and beyond
297 We were concerned that our conclusions might be biased by the particular populations we 298 happened to sample, and that we would have obtained qualitatively different conclusions 299 without data from some key populations. We tested our conclusions by plotting the inferred 300 position of admixed populations in PCA against a weighted combination of their inferred 301 source populations and obtained qualitatively consistent results [Extended Data Fig. 5].
302 To further assess the robustness of our inferences, we developed a method to infer the 303 existence and genetic affinities of ancient populations from unobserved ‘ghost’ populations 304 [Supplementary Information, section 10; Extended Data Fig. 6]. This method takes advantage 305 of the insight that if an unsampled ghost population admixes with differentiated ‘substratum’
9
bioRxiv preprint first posted online Jun. 16, 2016; doi: http://dx.doi.org/10.1101/059311. The copyright holder for this preprint [which was not peer-reviewed] is the author/funder. It is made available under a CC-BY-NC-ND 4.0 International license.
306 populations, it is possible to extrapolate its identity by intersecting clines of populations with 307 variable proportions of ‘ghost’ and ‘substratum’ ancestry. Applying this while withholding 308 major populations, we validated some of our key inferences, successfully inferring mixture 309 proportions consistent with those obtained when the populations are included in the analysis. 310 Application of this methods highlights the impact of Ancient North Eurasian [ANE] ancestry 311 related to the ~22,000 BCE Mal’ta 1 and ~15,000 BCE Afontova Gora 215 on populations 312 living in Europe, the Americas, and Eastern Eurasia. Eastern Eurasians can be modelled as 313 arrayed along a cline with different proportions of ANE ancestry [Supplementary
314 Information, section 11; Extended Data Fig. 7], ranging from ~40% ANE in Native 315 Americans matching previous findings13,15, to no less than ~5-10% ANE in diverse East 316 Asian groups including Han Chinese [Extended Data Fig. 4; Extended Data Fig. 6f]. We also 317 document a cline of ANE ancestry across the east-west extent of Eurasia. Eastern Hunter 318 Gatherers [EHG] derive ~3/4 of their ancestry from the ANE [Supplementary Information, 319 section 11]; Scandinavian hunter-gatherers7,8,13 [SHG] are a mix of EHG and WHG; and 320 WHG are a mix of EHG and the Upper Paleolithic Bichon from Switzerland [Supplementary 321 Information, section 7]. Northwest Anatolians—with ancestry from a population related to 322 European hunter-gatherers [Supplementary Information, section 7]—are better modelled if 323 this ancestry is taken as more extreme than Bichon [Supplementary Information, section 10].
324 The population structure of the ancient Near East was not independent of that of Europe 325 [Supplementary Information, section 4], as evidenced by the highly significant [Z=-8.9] 326 statistic f4[Iran_N, Natufian;WHG, EHG] which suggests gene flow in ‘northeastern’ 327 [Neolithic Iran/EHG] and ‘southwestern’ [Levant/WHG] interaction spheres [Fig. 4d]. This 328 interdependence of the ancestry of Europe and the Near East may have been mediated by 329 unsampled geographically intermediate populations37 that contribute ancestry to both regions.
330 Conclusions
331 By analysing genome-wide ancient DNA data from ancient individuals from the Levant, 332 Anatolia, the southern Caucasus and Iran, we have provided a first glimpse of the 333 demographic structure of the human populations that transitioned to farming. We reject the 334 hypothesis that the spread of agriculture in the Near East was achieved by the dispersal of a 335 single farming population displacing the hunter-gatherers they encountered. Instead, the 336 spread of ideas and farming technology moved faster than the spread of people, as we can 337 determine from the fact that the population structure of the Near East was maintained
10
bioRxiv preprint first posted online Jun. 16, 2016; doi: http://dx.doi.org/10.1101/059311. The copyright holder for this preprint [which was not peer-reviewed] is the author/funder. It is made available under a CC-BY-NC-ND 4.0 International license.
338 throughout the transition to agriculture. A priority for future ancient DNA studies should be 339 to obtain data from older periods, which would reveal the deeper origins of the population 340 structure in the Near East. It will also be important to obtain data from the ancient 341 civilizations of the Near East to bridge the gap between the region’s prehistoric inhabitants 342 and those of the present.

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the lioness,
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http://biorxiv.org/content/early/2016/06/16/059311

Iosif Lazaridis, comments:

By including Natufians into the Right set of outgroups we are able to distinguish between Levantine and other populations, as Natufians share more alleles with Levantine Neolithic than with other ancient populations.

qpAdm does not really make use or depend on admixture time. The data is consistent with East Africans' allele sharing with Natufians being mediated by Levantine Neolithic. This does not mean that the admixture took place before, during, or after the Levantine Neolithic, only that the best source for it we could find was Levant_N, which may of course have existed prior, during, and after the time of our particular samples.

Analysis of present-day populations cited in our paper suggest a fairly recent admixture even [~3,000 years]. The fact that Mota did not yet have it also suggest this [although in principle Mota could be an isolated population that did not experience it]. Of course, it will be nice to see this directly with ancient DNA from East Africa.

To illustrate what I mean about possible older dates, I would point to southern Africans. Recent studies have similarly found and dated admixture in southern Africa to <2 kya. If we did not have a modern proxy for their West Eurasian ancestry [East African pastoralists], one may have assumed that there was a Eurasian migration into southern Africa at that time. Yet, we know this is not true; rather, it is the date of population intermixture, with East Africans of partial Eurasian ancestry.

Although it was not the focus of your rather comprehensive study,
the simple model originally proposed by Pagani 2012, whereby all of the
Eurasian ancestry in East Africa derives from the known Semitic
migration, has not been adequately scrutinized in my opinion.

The reconstructed proto-Cushitic language, which predates the 3 kya timeframe by thousands of years, had words for sheep and goats. These words denote a pastoral lifestyle, perhaps imported from the PPNB culture which you have sampled. Mota, a hunter-gatherer living in a remote part of Ethiopia, is not necessarily representative of early pastoral Africans living closer to the Nile Valley.

Sheep and goats appeared in NE Africa after the appearance of the Levantine PPNB culture, and in light of this, your paper's results make a lot of sense, even if there may have been later intermixture ~3 kya which muddled the original admixture event. After all, LD-based admixture dating methods are known to be biased toward the most recent admixture event.

I accept not having Natufians among tested populations in this particular qpAdm analysis. Other comparisons may be informative however, particularly considering the relative African Fst shift noted in a different comment.

[ I1290 is genetically a female as stated in Supplementary Data Table 1 and not a male; we will correct SI1 in the revision.]

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LINK

http://biorxiv.org/content/biorxiv/suppl/2016/06/16/059311.DC1/059311-1.pdf


_______________________

wikipedia:

E-Z830 (E1b1b1b2)

This is a recently discovered subclade which has not yet been included in most haplogroup trees, E-Z830 includes the confirmed subclades of E-M123, E-V1515 (E-M293, E-V42, E-V6, E-V92), and E-Z830*, and is a sibling clade to E-L19. Currently, the E-M35 phylogeny project recognizes four distinct clusters of Z830* carriers, two of which are exclusively Jewish in origin. The remaining two are significantly smaller, and include scattered individuals in Germany, Spain, Latin America, Egypt, and Ethiopia.

source:

http://www.haplozone.net/e3b/project/cluster/3

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xyyman
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Are you saying Jews are black Africans?

--------------------
Without data you are just another person with an opinion - Deming

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quote:
Originally posted by xyyman:
Are you saying Jews are black Africans?

I didn't say anything about black. Why are you bringing race into this?
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xyyman
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1. “Black” is not a race since there is no race. Black skin exist all over the planet, not only in Africa similar for “Negroids”. Exist all over the planet. Modern Africans do NOT have monopoly on Black skin or negroid.
2. Jews undoubtedly have a strong African influence. If you are R1b or mtDNA H1/H3 etc you are NOT a Jew. The Jew heritage may carry E-M35 or J1-M267 and mtDNA L mostly.

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