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Sunday, July 16, 2017

North European admixture in the Han Chinese (Charleston et al. 2017 preprint)

Over at bioRxiv at this link. Emphasis is mine. The estimated date of the North European-related admixture signal is probably much too late. These sorts of estimates always look way off. And I doubt that it's largely the result of the Silk Road, which linked China to the Near East and Mediterranean rather than to Northern Europe. More likely it reflects gene flow from the Pontic-Caspian steppe in Eastern Europe during the Bronze and Iron ages, via the Afanasievo, Andronovo, and other closely related steppe peoples (see here).

Abstract: As are most non-European populations around the globe, the Han Chinese are relatively understudied in population and medical genetics studies. From low-coverage whole-genome sequencing of 11,670 Han Chinese women we present a catalog of 25,057,223 variants, including 548,401 novel variants that are seen at least 10 times in our dataset. Individuals from our study come from 19 out of 22 provinces across China, allowing us to study population structure, genetic ancestry, and local adaptation in Han Chinese. We identify previously unrecognized population structure along the East-West axis of China and report unique signals of admixture across geographical space, such as European influences among the Northwestern provinces of China. Finally, we identified a number of highly differentiated loci, indicative of local adaptation in the Han Chinese. In particular, we detected extreme differentiation among the Han Chinese at MTHFR, ADH7, and FADS loci, suggesting that these loci may not be specifically selected in Tibetan and Inuit populations as previously suggested. On the other hand, we find that Neandertal ancestry does not vary significantly across the provinces, consistent with admixture prior to the dispersal of modern Han Chinese. Furthermore, contrary to a previous report, Neandertal ancestry does not explain a significant amount of heritability in depression. Our findings provide the largest genetic data set so far made available for Han Chinese and provide insights into the history and population structure of the world's largest ethnic group.


One finding from our analysis of admixture signals that most likely fit a one-pulse admixture model is our observation of admixture from Northern European populations to the Northwestern provinces of China (Gansu, Shaanxi, Shanxi), but not other parts of China. Previous analysis of the HGDP data, based on patterns of haplotype sharing among 10 Han Chinese from Northern China, estimated a single pulse of ~6% West Eurasian ancestry among the Northern Han Chinese. The estimated date of admixture was around 1200 CE. This signal is also observed among the Tu people, an ethnic minority also from Northwestern China; the authors attributed this signal to contact through the Silk Road (Hellenthal et al. 2014). We estimate a lower bound of admixture proportion due to Northern Europeans at approximately 2%-5%, with an admixture date of about 26 +/-3 generations for Gansu, and 47 +/-3 generations for Shaanxi [Table S8]. Using a generation time of about 26-30 years (Moorjani et al. 2016), these estimates correspond to admixture events occurring at around 700 CE and 1300 CE, respectively, corresponding roughly to the Tang and Yuan dynasty in China. However, these estimated dates should be interpreted with caution, as both the violation of a single pulse admixture model and the additional noise in inter-­marker LD estimates due to low coverage data could bias the estimates.

Charleston et al.,A comprehensive map of genetic variation in the world's largest ethnic group - Han Chinese, bioRxiv, Posted July 13, 2017, doi:

See also...

Late PIE ground zero now obvious; location of PIE homeland still uncertain, but...


Ryan said...

~47 generations ago would line up with when the Jie disappeared from Shanxi.

MaxT said...

"our observation of admixture from Northern European populations to the Northwestern provinces of China (Gansu, Shaanxi, Shanxi), but not other parts of China."

Northwestern provinces...sounds like steppe-like admixture. I would guess it's Andronovo or even Scythians.

MaxT said...

First appearance of horses and chariot in China during Shang Dynasty, some historians have recently claimed that Indo-Europeans were responsible for the foundation of the Shang Dynasty.

"The chariot first appeared in China around 1200 BC, during the reign of Wu Ding. There is little doubt that the chariot entered China through the Central Asia and the Northern Steppe, possibly indicating some form of contact with the Indo-Europeans. Recent archaeological finds have shown that the late Shang used horses, chariots, bows and practiced horse burials that are similar to the steppe peoples to the west. Christopher I. Beckwith speculates that Indo-Europeans may even have been responsible for the foundation of the Shang Dynasty".

MaxT said...

Shang dynasty geography fits well with the studies findings "Northwestern provinces of China (Gansu, Shaanxi, Shanxi) but not other parts of China". Looks like there could actually be some truth to historian Beckwith's claims!

MaxT said...

Davidski, can you test their northern Han samples to see what kinda of steppe admixture they carry, Steppe_EMBA or Steppe_MLBA?

mickeydodds1 said...

Curiouser and curiouser.

Is there anything the steppe/corded ware people did not do?

Davidski said...


I don't have access to their genomes, but the Tu people from NW China show Steppe MLBA type of admix, which is very close to modern N/E European.

MaxT said...

"but the Tu people from NW China show Steppe MLBA type of admix, which is very close to modern N/E European."

Thanks, Steppe_MLBA close to China would be Andronovo-related admixture, instead of older Afanasevo culture/Steppe_EMBA.

epoch2013 said...

Is this the source of the Han signal in the Villabruna cluster? That was consistently accompanied by a Karitiana signal, which could point to ANE or ANE related ancestry. If this indeed is the Villabruna signal, the Fu et al D-stats should show that signal for South-Western Chinese groups.

epoch2013 said...

O I really make a mess out this:

That should have read: "If this indeed is the Villabruna signal, the Fu et al D-stats should not show that signal for South-Eastern Chinese groups."

Ryan said...

I doubt it's the source of the Villabruna signal. Based on Y-chromosomes alone we can be pretty sure there's some sort of very deep East Asian influence there.

MaxT said...


No, the study gives admixture events only between 7th A.D to 13th century A.D

"these estimates correspond to admixture events occurring at around 700 CE and 1300 CE, respectively, corresponding roughly to the Tang and Yuan dynasty in China."

It's possible some admixture events among northern Han dates further back to Shang, when horses/chariots first appeared in China.

epoch2013 said...


What does this do?
Kostenki14 Loschbour Han Mbuti
Vestonice16 Loschbour Han Mbuti
Kostenki14 Loschbour Dai Mbuti
Vestonice16 Loschbour Dai Mbuti
Kostenki14 Iberia_Mesolithic Han Mbuti
Vestonice16 Iberia_Mesolithic Han Mbuti
Kostenki14 Iberia_Mesolithic Dai Mbuti
Vestonice16 Iberia_Mesolithic Dai Mbuti

epoch2013 said...


But although the admixture event maybe fairly recent, the admixture brought into Han could be related to Villabruna.

Davidski said...

result: Kostenki14 Loschbour Han Mbuti -0.0182 -3.636 22556 23391 476035
result: Vestonice16 Loschbour Han Mbuti -0.0118 -2.319 17286 17700 373683
result: Kostenki14 Loschbour Dai Mbuti -0.0157 -3.033 22626 23347 476035
result: Vestonice16 Loschbour Dai Mbuti -0.0116 -2.194 17313 17721 373683
result: Kostenki14 Iberia_Mesolithic Han Mbuti -0.0215 -4.513 23936 24988 502541
result: Vestonice16 Iberia_Mesolithic Han Mbuti -0.0171 -3.349 17782 18400 379342
result: Kostenki14 Iberia_Mesolithic Dai Mbuti -0.0198 -4.050 23976 24947 502541
result: Vestonice16 Iberia_Mesolithic Dai Mbuti -0.0170 -3.199 17787 18401 379342

Davidski said...

The North European admixture could be related to the rise of the Shang Dynasty. See here.

The origin of Chinese domestic horses revealed with novel mtDNA variants

EastPole said...

Chinese use Slavic word ‘kolo’ for ‘wheel’ which is interesting:

Also Chinese ‘mjit/mit’ for honey is very interesting:

Shaikorth said...


Definitely steppe admixture. With those dates and a northwestern distribution the population mixing with Han could've been eastern Scythian, Tocharian or even some Turkic group with related ancestry. ALDER or f3 mixture test with just modern European references can't precisely tell which one of them.

Ebizur said...

Please keep in mind that there are more than ten million Chinese-speaking Muslims (Hui people) in China, and they clearly contain a lot of exotic admixture (at least in their Y-DNA):

Hui (Xue et al. 2006)
2/35 = 5.7% BT-SRY10831.1(xC-M130, DE-YAP, J-12f2, K-M9)
4/35 = 11.4% C-M130(xM8, M217)
4/35 = 11.4% C-M217(xM48)
4/35 = 11.4% DE-YAP(xE-M40)
1/35 = 2.9% E-M40
3/35 = 8.6% J-12f2
1/35 = 2.9% NO-M214(xM175, M128, P43, Tat)
3/35 = 8.6% O-M119
1/35 = 2.9% O-P31(xM95, M176)
4/35 = 11.4% O-M122(xM159, M7, M134)
1/35 = 2.9% O-M134(xM117)
1/35 = 2.9% O-M117
4/35 = 11.4% P-92R7(xR1a1-SRY10831.2)
2/35 = 5.7% R1a1-SRY10831.2

Hui in Yunnan (Zhong et al. 2011)
1/10 C-M217
1/10 J1-M267
2/10 N-M231
3/10 O-M175
1/10 Q-M346
1/10 R-M17
1/10 R-M335

Hui in Ningxia (Zhong et al. 2011)
2/62 = 3.2% D-M174
2/62 = 3.2% E-M40
5/62 = 8.1% C-M130(xM217, M356)
6/62 = 9.7% C2-M217(xM93, P39, M48, M407)
1/62 = 1.6% G1-M285
1/62 = 1.6% G2a-P15
1/62 = 1.6% H1a-M69/M370(xH1a1-M52)
1/62 = 1.6% J1-M267
5/62 = 8.1% J2a-M410(xM322, M67, M319)
2/62 = 3.2% J2a1b-M67
1/62 = 1.6% L1a2-M357
5/62 = 8.1% N-M231
22/62 = 35.5% O-M175
2/62 = 3.2% Q1a1a-M120
1/62 = 1.6% R1-M173(xR1a1a-M17, R1b-M343)
2/62 = 3.2% R1a1a-M17
1/62 = 1.6% R1b1a1a2-M269
2/62 = 3.2% R2a-M124

However, R1a1-SRY10831.2 has been found in samples of non-Muslim Han in Northwest China, too:

Han in Lanzhou, Gansu (Xue et al. 2006)
6/30 = 20.0% C-M217(xM48)
2/30 = 6.7% DE-YAP(xE-M40)
3/30 = 10.0% J-12f2
2/30 = 6.7% NO-M214(xM175, M128, P43, Tat)
1/30 = 3.3% O-M175(xM119, P31, M122)
2/30 = 6.7% O-M119
1/30 = 3.3% O-M95(xM88)
6/30 = 20.0% O-M122(xM159, M7, M134)
2/30 = 6.7% O-M134(xM117)
3/30 = 10.0% O-M117
2/30 = 6.7% R1a1-SRY10831.2

Han in Yili, Xinjiang (Xue et al. 2006)
1/32 = 3.1% BT-SRY10831.1(xC-M130, DE-YAP, J-12f2, K-M9)
2/32 = 6.3% C-M217(xM48)
1/32 = 3.1% DE-YAP(xE-M40)
3/32 = 9.4% K-M9(xNO-M214, P-92R7)
3/32 = 9.4% O-M119
3/32 = 9.4% O-P31(xM95, M176)
1/32 = 3.1% O-M95(xM88)
10/32 = 31.3% O-M122(xM159, M7, M134)
2/32 = 6.3% O-M7
1/32 = 3.1% O-M134(xM117)
2/32 = 6.3% O-M117
1/32 = 3.1% P-92R7(xR1a1-SRY10831.2)
2/32 = 6.3% R1a1-SRY10831.2
(Has 92R7 been removed from the tree because of unreliability? Perhaps 3/32 K-M9(xNO-M214, P-92R7) and 1/32 P-92R7(xR1a1-SRY10831.2) should be combined as 4/32 K-M9(xNO-M214, R1a1-SRY10831.2).)

I suspect that Chinese speakers in Northwest China have multiple layers of non-East Asian ancestry.

Ric Hern said...

Wonder where that BT came from ?

Salden said...

I suspect the PRC progapanda arm wouldn't be pleased about this.

Ebizur said...

"Wonder where that BT came from ?"

It almost certainly is some sort of G, H, I, or F* (e.g. F2-M427/M428) that simply has not been tested for the relevant SNPs.

wagg said...

There are tracks of a migration from the Russian steppes to the east at the end of neolithic/early Bronze age (Afanasevo, which seems related in many ways to early Yamnaya culture) and these ancient tracks are seen at least up to the Baikal Lake.

"From the Altay-Sayan steppes the Europeoid groups seem to have moved fairly far east; the neolithic population west of Lake Baykal, in particular, shows a Europeoid admixture." ((M. G. Levin, “The Anthropological Types of Siberia,” in The Peoples of Siberia, ed. M. G. Levin and L. P. Potapov, The University of Chicago Press, 1964 - Page 99)).

These populations are likely at the origin of the later Tocharian languages - though it's unproved - but these like proto-Indo-European speakers (given the timeframe) might have not all ended up in south Siberia and Xinjiang, some might have gone further, going maybe up to south-eastern Siberia and/or in north China, where they were eventually absorbed without leaving much traces.

The fact that some chinese words could be of IE origins (some Tocharians, but some could be derived from PIE) give some weight to it (but there are no real prooves).
Quan (PIE *kwon) = dog
Ku (Tocharian ku) = dog
Mi (Tocharian mid) = honey (this root for honey is found everywhere in the IE world from Old Irish (mid) to Sanskrit (madhu)).
Yang (from an older [g]riang that could be linked with Tokharien yriye (“lamb”)).
There's also a dialectal Chinese word gulu (wheel) from an ancient form *kolo that reminds of Slavic kolo and old Prussian kelan.

There are also a resemblances between Mandarin Nyu (=cow) from reconstructed Old Chinese (nguw, maybe (see also Cantonese Ngau and Hakka ngiu)) and PIE *gwou-. The problem is that the stem seems to pertain to an ancestral Proto-Sino-Tibetan word reconstructed as ngwa, so maybe it's just a coincidence. Unless they all pertain to a loanword from China (from IE dialect) spreading with cattle but that doesn't seem the most likely.

And the impact might have gone further than China:

In "Mitochondrial DNA variation and evolution of Japanese black cattle (Bos taurus)" ( it is written "Unlike Africa, half of Japanese cattle sequences are topologically intermingled with the European variants. This suggests an interchange of variants that may be ancient, perhaps a legacy of the first introduction of domesticates to East Asia".

Add to that the Ainu (an ancient autochtonous Japanese ethnic group) and some Japanese dialect words for “cattle” and “cow” are very similar to the Proto-indo-european root *peku- (e.g. latin pecus, sanskrit pasu, gothic faihu, etc...) : peko (Ainu) and beko (in some Japanese dialects)

The Mongolian cattle was also shown to also have western genes, showing a trail from the west going deep in south siberia.

wagg said...

Notice also that in "Genetic Structure of a 2,500-Year-Old Human Population in China and Its Spatiotemporal Changes" ( it was found that "the 2,500-year-old Linzi population showed greater genetic similarityto present-day European populations than to present-day east Asian populations. The 2,000-year-old Linzi population had features that were intermediate between the present-day European/2,500-year-old Linzi populations and the present-day east Asian populations. These relationships suggest the occurrence of drastic spatiotemporal changes in the genetic structure of Chinese people during thepast 2,500 years.”.


this was verified in "Reanalysis of Eurasian Population History: Ancient DNA Evidence of Population Affinities (Bennett, Casey C., Kaestle, Frederika A. | Human Biology – Volume 78, Number 4, August 2006, pp. 413-440) (;col1) where it is said that "The Linzi material seems to bear a stronger affinity with Near Easterners and Europeans rather than with the present-day populations of northern China, although there is a definite component of East and/or Southeast Asians within Linzi as well"

wagg said...

BTW, concerning the Japanese bit, I'm not saying some IE speakers reached Japan of course, just that the populations that migrated in Japan a few centuries BCE (likely from south-eastern Siberia and creating the Yayoi culture IIRC) got this loanwaord with some of its cattle likely in south Siberia from people with at least partly some IE origins (and the loanword eventually reached the ainus fomr these newcomers on the island).

epoch2013 said...


What surprises me is that the outcome of these is different from the supp info in Fu et al.

K14 Loschbour; Han Mbuti Z=-4.3
Vestonice16 Loschbour; Han Mbuti Z=-3.9
K14 LaBrana1; Han Mbuti Z=-3.8
Vestonice16 Loschbour; Han Mbuti Z=-3.4

Anyway, never mind. There seems to be not much difference between Han and Dai, a Yunnan minority from South-East China.

Ebizur said...

Japanese (dialect) beko is probably of onomatopoetic origin, and was borrowed by the Ainu after the introduction of cattle into Hokkaido by Japanese settlers. (By the way, the Ainus' northern neighbors, the Nivkhs, borrowed the Tungusic appellation for the animal: cf. Nivkh eɣa(ŋ) and Manchu ihan).

Beko or beeko meaning "bovine" is attested in Japanese dialects of Hokkaido and Tohoku (Sendai, Nambu, Akita) since at least the Edo Period. It is still used today in various dialects in the Tohoku region and neighboring areas (some parts of Niigata and Tochigi prefectures) as well as far off to the southwest in the areas of Kyoto and Hyogo prefectures that face the Sea of Japan. In many other parts of central and western Honshu as well as Shikoku, it (or a similar word, bekko) means specifically "calf." There are also a few dialects, such as in part of Kyushu, where the word means specifically "cow, female bovine." The shorter form bee is used in many parts of Kyushu and some parts of Honshu (mainly along the Pacific coast of central Honshu) to mean "calf." All these forms probably derive from an onomatopoetic root attested in Standard Japanese as mee, which is used to mimic the cry of a goat (and sometimes other animals, such as sheep). The speakers of some dialects use mee either to mimic the cry of a cow or as a way to call a cow toward oneself; speakers of Standard Japanese would say moo instead. Voiced oral plosives such as /b/ do not normally occur in word-initial position in Japanese words, which strongly suggests either an onomatopoetic or a foreign origin of this group of dialect words; I strongly support the hypothesis of an onomatopoetic origin (especially because of the Kyushu/Honshu Pacific coast form /bee/ "calf").

Grey said...

mickeydodds1 said...
"Is there anything the steppe/corded ware people did not do?"

I think there's a certain inevitability to some aspects of this. If you divide land into two groups a) suitable for crop farming and b) too marginal for crop farming resulting in a farmer population living on a) and a herder population living on b) and assume conflict will cyclically occur then if the farmers win they still can't move onto the herder's land - they might kill a lot of herders but the population eventually regrows. On the other hand if the herders win they can move onto the farmer's land and either displace them (if the farmer's population density is low) or become a ruling caste (if the farmer's population density has become too high). So the farmers have to win every periodic conflict while the herders only have to win once.

Some exceptions to this deterministic rule might be:
1) some physical barrier that constrains the herders like mountains or a narrow river valley between mountains which is undrained and therefore still a giant swamp
2) the farmers improving their techniques to the point where they have too big a numerical advantage
3) the farmers improving their techniques to the point where they can expand over previously marginal land thus denying herders living space
4) guns

(having said that my guess is this admixture could also be infiltration migration - steppe herders moving onto land that was too marginal for crop farming but good for cattle (or sheep, goats etc))

Ric Hern said...

Thanks Ebizur.

Ric Hern said...

@ wagg

Interesting. One of my friends have an interesting theory that shows many connections between Chinese and Q-Celtic...too many to be pure coincidence.

Matt said...

New paper (probably basis for next Eurogenes post I'd guess) : - "Genomics of Mesolithic Scandinavia reveal colonization routes and high-latitude adaptation"

Peter Klevius said...

'kolo' is a Finnish word still in use and meaning 'hole'. All Uralic languages have it in slightly varying forms and its stem is proposed as *kol3. Slavic has borrowed it more recently.

Salden said...

>Such results suggest that the blue eye-color allele is rather old. Using an ABC modeling
approach Nakagome et al. (186), predicted that the light-pigmentation allele at rs12913832
emerged around 42,000 years ago or earlier; a date close in time to the initial peopling of Europe.
>A plausible scenario of the origin of the blue-eye mutation that reconciles our results with
findings from other studies is one where this variant appeared in an ancestral population before
the ancestors of the WHG migrated from Near East into West and Central Europe.

From the Scandinavia Paper. This is interesting when you consider how light eye colors are prominent in MENA minorities like select Berbers and Christians. Both of which are isolated or endogamous.

JohnP said...

The 3 godly peoples of pre-history in my opinion were the First Farmers, Indo-Europeans and the people of the Elam-Sumeria-Egypt area.
They were vital to humanity, or at least to West Eurasia, but then we see things now and then saying that Indo-Europeans gave East Asians many technologies, etc and then you start wondering if they really had something special, those orange, teal and blue.

Strandloper said...

Some day we will find that they did not stop in china but made all the way to the Americas to influence the civilizations there as well.

europeuplement said...

"these estimates correspond to admixture events occurring at around 700 CE and 1300 CE, respectively, corresponding roughly to the Tang and Yuan dynasty in China."

possibly Sogdians (Sogdian dérived from Scythian people)

previously, Ordos culture (a scythian culture), settled in China

Slumbery said...

The Wikipedia article you linked states the results were errors and there is no particular Western affinity in 2500 y BP Linzi. Indeed it would be super surprising if the population of the capital of an Eastern Chinese kingdom came out as genetically super Western.

EastPole said...

@Peter Klevius
“'kolo' is a Finnish word still in use and meaning 'hole'. All Uralic languages have it in slightly varying forms and its stem is proposed as *kol3. Slavic has borrowed it more recently.”

I don’t think Slavs borrowed it from Uralic speakers recently. Slavs originated from Corded Ware in the area where first wheeled vehicles were used:

Possible links between Uralic and Slavic languages can be also explained by common origin.
Balto-Slavs originated on the steppe before 3000 BC and as a steppe population they were a mixture of EHG and Caucasus farmers. As Uralic speaker are also partially descendent from EHG our common words can have their origin in EHG language.

Slavs originated after 3000 BC when steppe Balto-Slavic population started to mix with Anatolian farmers in Western Ukraine/Poland area and as Corded Ware expanded with wheeled vehicles which is confirmed by the distribution of R1a-Z645 and the presence of Anatolian farmers component in Sithashta and Andronovo.

In addition to ‘kolo’ which can mean big wheel or an area around we also have a diminutive ‘kulko’ which means small wheel and can be applied to wheeled vehicles as well. After metathesis ‘kulko’ > ‘kuklo’ this word has been spread to other IE languages.

Lukasz M said...

Northwestern provinces...sounds like steppe-like admixture. I would guess it's Andronovo or even Scythians.

I guess Tocharians.

P Piranha said...

Getting waaaay off base here.

Beckwith's idea that the Shang, and Chinese Bronze Age civilisation in general, was founded by IEs is a very fringe one, with very little evidence to support it. IE loanwords in old Chinese are present, however they are concentrated in certain peripheral cultural fields relating to Chariots, Wool, etc and the language of both early Chinese dynasties, which is recorded in writing, is clearly Sinitic. Early Chinese spirituality and burial customs have clearly almost nothing to do with steppic traditions--ancestor-worship is almost always found among agriculturalists and is not important in Steppic cultural worlds--and the same for Chinese metallurgy, which is very original and idiosyncratic, so much that Southeast Asian metallurgy resembles steppic models more than Chinese metallurgy does. The earliest state-like societies in the Yellow River Valley in China seem to emerge from influences from the South and Southwest, not the North, judging from stable elements of culture such as burial rites and supernatural conceptions.

"Nordic/Europoid" and "Pamirid" skulls were among those sacrificed in Shang tombs, clearly placing them as a class of foreign enemies. Its likely that the early IEs in China played an analogous role to those in early Mesopotamia, with tribes or mercenaries hired into what were already highly complex societies in the lowlands.

IE influence, and just steppic influence in general, on the second major Chinese Bronze Age dynasty, the Zhou, is much clearer, which is expected as they did emerge from the Northwest; the Zhou do have a much more prominent "high-moralising God" similar to that found on the Steppes in addition to ancestor-worship as an additional basis for spirituality for example, but that is more similar to Altaics than IEs and is also well into the historical period.

wagg said...

@Slumbery:"The Wikipedia article you linked states the results were errors and there is no particular Western affinity in 2500 y BP Linzi"

Yes, I know.
What the "author" of this wikipedia page ( conveniently forgot to say is that there was a new analysis done in 2006 that supported the results of the first one in 2000 (IIRC).
The second part of my post you're commenting was from the second DNA analysis done in 2006 (while the Chinese rebuttal was done in 2003).

Unknown said...

>From the Scandinavia Paper. This is interesting when you consider how light eye colors are prominent in MENA minorities like select Berbers and Christians. Both of which are isolated or endogamous.

Berbers have high SSA admix so not very isolated or endogamous.