Revised timescale of human mtDNA evolution (Fu et al. 2013)

 

An important new paper has just appeared in Current Biology. It is very exciting for a couple of reasons:

• The paper uses the idea of branch shortening to infer dates for the mtDNA phylogeny. Briefly, if one counts differences between a present-day sample P and an ancient sample A, one can get a sense of when their most recent common ancestor X, lived. But PX and PA are not equal: PA is shorter, because A has "missed" a few thousand years of evolution (depending on its age). If we know the age of A (and this can be reliably known by direct dating for many samples), then we can infer the time of X.

Looking at the ages of many haplogroups, I don't see any that immediately strike me as inconsistent with recent published age estimates based on modern mtDNA alone, such as those from the Copernican reassessment paper. In any case, this is an exciting new application of an idea that will yield good dates for haplogroup ages in the future. For example, it could be used to date Y-chromosome lineages as well, when high-quality sequences of ancient human Y-chromosomes become available.

• The paper also presents a number of new and exciting ancient DNA samples:

Please note that the "Cro Magnon" sample is actually revealed to be non-ancient in the study; that is another utility of the "branch shortening" idea, since it can demonstrate that purported very old samples are in fact fairly recent.
Note the important new samples from Dolni Vestonice, Oberkassel, Continenza, all of which are Paleolithic Europeans and all of which belong, without exception to subgroups of haplogroup U. It appears that Europe was indeed dominated by this haplogroup down to Mesolithic times, with a sharp discontinuity with early Neolithic Europeans.
The only outlier in the European context is Paglicci Str. 4b which is assigned to haplogroup H1. However:

Using this criterion, we excluded Paglicci Str. 4b from further analysis as the rate of C to T misincorporation at the 50 end was only 8.8%, thus making an ancient origin for the DNA in this sample uncertain [14].

Personally, I doubt there was any mtDNA haplogroup H in pre-Neolithic Europe, as the first author of this paper has also argued for in a previous one.
There is also a new sample from Boshan; it is probably too early to detect a pattern, but it is nonetheless noteworthy that it also belonged to mtDNA haplogroup B like the much earlier sample from Tianyuan.
Finally, the paper also discusses the issue of the mutation rate, noting that the divergence between Eurasian (M+N) and African (L3) mtDNA is much later than that inferred for autosomal DNA using new and "slow" de novo autosomal mutation rates. In my opinion there are two possible interpretations for this: one that there's something wrong with the slow rates, while another is that the earlier divergence using autosomal DNA may be a consequence of inflation due to admixture events in Africa.
Current Biology doi:10.1016/j.cub.2013.02.044
A Revised Timescale for Human Evolution Based on Ancient Mitochondrial Genomes
Qiaomei Fu et al.
Summary
Background
Recent analyses of de novo DNA mutations in modern humans have suggested a nuclear substitution rate that is approximately half that of previous estimates based on fossil calibration. This result has led to suggestions that major events in human evolution occurred far earlier than previously thought.
Results
Here, we use mitochondrial genome sequences from ten securely dated ancient modern humans spanning 40,000 years as calibration points for the mitochondrial clock, thus yielding a direct estimate of the mitochondrial substitution rate. Our clock yields mitochondrial divergence times that are in agreement with earlier estimates based on calibration points derived from either fossils or archaeological material. In particular, our results imply a separation of non-Africans from the most closely related sub-Saharan African mitochondrial DNAs (haplogroup L3) that occurred less than 62–95 kya.
Conclusions
Though single loci like mitochondrial DNA (mtDNA) can only provide biased estimates of population divergence times, they can provide valid upper bounds. Our results exclude most of the older dates for African and non-African population divergences recently suggested by de novo mutation rate estimates in the nuclear genome.
Link

Doubt cast on Tam Pa Ling age

For reference, see the post on the original article:

• Tam Pa Ling: modern humans in Southeast Asia at 63-46ka

From a criticism of the original paper:

Hence, contrary to the authors' assertion that TPL1 has a “minimum secured age of 46 ka and a maximum age of ∼63 ka”, the published stratigraphy, if correct, indicates that the TPL1 specimen is no older than 46 ka. TPL1’s status as “the earliest well-dated modern human fossil east of the Jordan Valley” also appears weak compared with the Liujiang specimen dated to ∼153 ka (3), the Callao Cave fossil in the Philippines dated to 67 ka (4), and, above all, the ∼100 ka modern fossil from Zhirendong (5), discovered only 484 km northeast of Tam Pa Ling in Southern China.

Irreconcilable differences between stratigraphy and direct dating cast doubts upon the status of Tam Pa Ling fossil

Alain Pierret et al.

Link

... and from a reply to the criticism by the authors:

They question the validity of the dating because of an apparent “reverse stratigraphy” (Fig. 1), whereby “older” dates are located higher in the section [i.e., 51.4 (14C) at 2.1 m] and “younger” dates are at the bottom of the section [i.e., 48 ka (optically stimulated luminescence [OSL]) and >49.2 ka (14C) at 4.3 m] (2). This criticism ignores the presented SEs (table 2 and table S1 in ref. 1), which make the results statistically equivalent (Table 1). More importantly, they ignore that the radiocarbon results are well beyond the accepted radiocarbon barrier of ∼40 ka (3), indicating that the charcoal has a minimum age of ∼40 ka. Bearing in mind these problems, we have conservatively estimated the burial age to be ∼46 ka according to the luminescence dating of the sediments. As the luminescence results are stratigraphically consistent, we perceive no irreconcilable differences between the stratigraphy and dating.

...

Finally, we interpret TPL1 as the earliest human fossil that is both well-dated and fully modern in morphology. Zhirendong demonstrates a mixture of archaic and modern traits, making it significant but not fully modern in appearance (4). Similarly, the metatarsal from Callao Cave is only diagnostic to the genus Homo given that it falls within “the morphological and size ranges of Homo habilis and H. floresiensis” (ref. 5, p. 123). Although the modernity of the Liujiang fossil is not questioned, it has no direct date and no secure stratigraphic provenance. It has been variably dated to ca. 20 ka, ca. 67 ka, 111 to 139 ka, and >153 ka (6), and this uncertain stratigraphic context has prevented many scholars from accepting any of the dates currently attributed to it (6).

Reply to Pierret et al.: Stratigraphic and dating consistency reinforces the status of Tam Pa Ling fossil

Fabrice Demeter et al.

Link

The case for earlier Out-of-Africa (Boivin et al. 2013)

The case for earlier Out-of-Africa (Boivin et al. 2013)

An informative review critical of the ~60kya coastal-Out-of-Africa hypothesis. On the left, the authors' estimate of the distribution of hominin groups during MIS5.

From the paper:

Another under-appreciated issue is the anomalous nature of the genetic evidence for a rapid spread of modern humans from Africa to Asia. Echoing the fossil date anomaly, the mtDNA branch lengths for sampled populations are longest for those which are farthest east, in Near Oceania, and shortest in the Asian areas that would have been encountered first (Merriwether et al., 2005; Oppenheimer, 2009). The real problem, however, is that the variation in branch lengths suggests that a single genotype engaged in the expansion actually existed for 30 ka, which does not support a rapid expansion. The anomaly can be explained by what we call an an ‘M buffer’ effect (see Supplementary material A) which implies that the branch ages we observe are considerable underestimates of the time of arrival of the genotype to these areas. Such anomalously long-lived genotypes have been directly observed through ancient DNA in species such as the Iberian lynx (Dalen et al., 2011).

and:

We have focused here on the possibility that the modern human exit recorded by fossil evidence in the Levant in MIS 5 does not represent a failed dispersal, and that in fact our species was not only in the Levant but also the Arabian peninsula during this marine isotope stage, and spread to India before the Toba eruption at 74 ka (Petraglia et al., 2007). Another valid hypothesis we do not explore here is that H. sapiens was able to leave Africa in MIS 6 via a grassland corridor (Frumkin et al., 2011; see also Scally and Durbin, 2012). Yet another is that our species dispersed out of Africa shortly after its first appearance c. 195 ka, in MIS 7 (Dennell and Roebroeks, 2005: 1102). One other possibility is that there were several, separate dispersals of our species out of Africa (Dennell and Petraglia, 2012). At the same time, we acknowledge that major demographic changes occurred in MIS 4 and MIS 3, perhaps explaining the relatively young mtDNA coalescence age in living populations. The increasing evidence for complexity as well as the clear patterns of bias for all records, whether archaeological, fossil or genetic, suggests the need for an open mind to multiple scenarios for Out of Africa, as well as for more rather than less complex models of H. sapiens dispersal across Eurasia.

Quaternary International doi:10.1016/j.quaint.2013.01.008

Human Dispersal Across Diverse Environments of Asia during the Upper Pleistocene

Nicole Boivin et al.

The initial out of Africa dispersal of H. sapiens, which saw anatomically modern humans reach the Levant in Marine Isotope Stage 5, is generally regarded as a ‘failed dispersal’. Fossil, archaeological and genetic findings are seen to converge around a consensus view that a single population of H. sapiens exited Africa sometime around 60 thousand years ago (ka), and rapidly reached Australia by following a coastal dispersal corridor. We challenge the notion that current evidence supports this straightforward model. We argue that the fossil and archaeological records are too incomplete, the coastal route too problematic, and recent genomic evidence too incompatible for researchers not to remain fully open to other hypotheses. We specifically explore the possibility of a sustained exit by anatomically modern humans, drawing in particular upon palaeoenvironmental data across southern Asia to demonstrate its feasibility. Current archaeological, genetic and fossil data are not incompatible with the model presented, and appear to increasingly favour a more complex out of Africa scenario involving multiple exits, varying terrestrial routes, a sub-divided African source population, slower progress to Australia, and a degree of interbreeding with archaic varieties of Homo.

Link

Genetic variation in gorillas quantified (Scally et al. 2013)

Genetic variation in gorillas quantified (Scally et al. 2013)

The allele frequency spectrum (Figure 3) from the paper is shown on the left. From the paper:

Figure 3 shows the resulting mean conditional AFS for nine western lowland gorillas (excluding the three lowest-coverage samples as above), and comparable samples from three human populations whose ancestry derives from Africa, Asia and Europe [22]. In a population whose effective size has remained constant, the theoretical expectation for such a conditional AFS is a straight line of constant negative slope [23], shown by the dashed line in Figure 3. Compared to this, western lowland gorillas show a deficit of rare alleles, consistent with their having undergone genetic drift due to a bottleneck or other reduction in effective population size during their demographic history. The similar signal in non-African human populations has been attributed to population contraction associated with the out-of-Africa event [24]. By contrast to the gorillas and the non-African humans, the African YRI population in Figure 3 shows an excess of rare alleles, consistent with population expansion and again similar to the signal seen in other African human data [24]. 

A population bottleneck culls rare alleles, and thus leads to a "dip" in the AFS on the left. An allele that occurs at, say, 10% frequency in a very small population is more likely to go extinct "by accident" than one which occurs at exactly the same frequency in a large population. You can think of this by taking the two extreme cases:

1. if an allele exists in exactly one physical copy (i.e., 1 copy in 5 diploid individuals = 10% frequency), then its bearer must survive, must reproduce, and the allele must be inherited by one offspring in order for it survive.
2. if the population has infinite size, then 10% frequency is still = infinite number of physical copies, hence the allele will survive no matter what [unless it always kills its unlucky bearer, but then how did it end up at 10% frequency in the first place?]

On the other hand, low-frequency alleles become more prevalent when the population expands in the recent past, because there is an ever growing number of bodies, an ever growing number of mutated alleles, but not enough time for these new alleles to grow in frequency.

A different mechanism whereby low-frequency alleles appear in excess in a population is admixture. African Americans, for example, have ~20% European ancestry, so any alleles that are present in Europeans and absent in Sub-Saharan Africans would tend to appear as low-frequency alleles in African Americans.

It is an open question to what degree modern human differences in the presence of low-frequency alleles are due to bottlenecks (such as the Out-of-Africa event) and to what degree they are due to admixture with other non-modern groups. A recent paper discovered a signal of Neandertal admixture when one considered alleles with 10% or less frequency in Europeans.

In the case of African Americans, we can tell that some of their low-frequency alleles were acquired by admixture with Europeans, because we have European samples; and, in the case of Europeans, we can tell that some of their low-frequency alleles were acquired by admixture with Neandertals, because we have a Neandertal genome. But, we don't have many of the genomes of archaic human groups that may have contributed variants to modern humans (and in some cases, like the Denisovans, we did not even know they existed in the first place!), so we must always keep in mind the possibility that such alleles may lurk on the left side of the AFS.

arXiv:1301.1729 [q-bio.PE]

A genome-wide survey of genetic variation in gorillas using reduced representation sequencing

Aylwyn Scally et al.

All non-human great apes are endangered in the wild, and it is therefore important to gain an understanding of their demography and genetic diversity. To date, however, genetic studies within these species have largely been confined to mitochondrial DNA and a small number of other loci. Here, we present a genome-wide survey of genetic variation in gorillas using a reduced representation sequencing approach, focusing on the two lowland subspecies. We identify 3,274,491 polymorphic sites in 14 individuals: 12 western lowland gorillas (Gorilla gorilla gorilla) and 2 eastern lowland gorillas (Gorilla beringei graueri). We find that the two species are genetically distinct, based on levels of heterozygosity and patterns of allele sharing. Focusing on the western lowland population, we observe evidence for population substructure, and a deficit of rare genetic variants suggesting a recent episode of population contraction. In western lowland gorillas, there is an elevation of variation towards telomeres and centromeres on the chromosomal scale. On a finer scale, we find substantial variation in genetic diversity, including a marked reduction close to the major histocompatibility locus, perhaps indicative of recent strong selection there. These findings suggest that despite their maintaining an overall level of genetic diversity equal to or greater than that of humans, population decline, perhaps associated with disease, has been a significant factor in recent and long-term pressures on wild gorilla populations.

Link

Introgression of archaic haplotype at OAS1 in Melanesians (Mendez et al. 2012)

Introgression of archaic haplotype at OAS1 in Melanesians (Mendez et al. 2012)

It seems that Michael Hammer was good on his promise that in 2012 "This year, we should be able to confirm what we found and go way beyond that."  In a new paper, conclusive evidence is presented about introgression of an archaic sequence into Melanesian populations. The argument is as follows:

• Melanesians are more diverse in that region than Africans.
• The common ancestor of the "archaic" and "African" haplotypes lived >3 million years ago.
• The "archaic" haplotype matches the ancient DNA from the Denisova hominin.
• Balancing selection (which can sometimes maintain extremely old polymorphism) is not reasonable in this case, because it would need to maintain both "archaic" and "African" haplotypes for a long time, but then (inexplicably) would continue to operate in Melanesia and cease to operate everywhere else.

Notice that once again, this is based on resequencing a small region of the genome. This is why I am all the more confident in my prediction that the advent of full genome sequencing will uncover more archaic admixture in humans. It may not always be able to use all the above listed criteria to confirm this admixture (since we do not and cannot have ancient DNA from all the archaic hominins that once roamed the planet), but all the remaining ones will suffice to make a very good case for introgression.

What I find particularly interesting, is that Mendez et al. re-iterate a few times that genomewide averages admit to different explanations:

Full genome comparisons of the Neandertal and Denisova draft genomes with modern human sequences have revealed different amounts of shared ancestry between each of these archaic forms and anatomically modern human (AMH) populations from different geographic regions. For example, a higher proportion of SNPs was shared between non-African and Neandertal, and between Melanesian and the Denisova genomes, than between either Neandertal or Denisova and extant African genomes (Green et al. 2010; Reich et al. 2010). An intriguing possibility is that these patterns result from introgression of archaic genes into AMH populations in Eurasia. However, this SNP sharing pattern could also be explained by ancestral population structure in Africa (i.e., without the need to posit introgression). For example, if non-Africans and the ancestors of Neandertals descend from the same deme in a subdivided African population, and this structure persisted with low levels of gene flow among African residents until the ancestors of non-Africans migrated into Eurasia, then we would expect more SNP sharing between non-Africans and Neandertals (Durand et al. 2011). 

... 

While genome-wide comparisons detect more sequence agreement between non-African and Neandertal genomes, and between Melanesian and Denisova genomes, the specific loci exhibiting these signals have not yet been identified. Furthermore, current analyses do not elucidate the relative roles of recent introgression versus long-term population structure in Africa in explaining these patterns.

The current paper does a good job at showing how in one particular region archaic introgression into Melanesians is indeed the best explanation for the evidence. But, the fact that the authors seem to re-iterate the possibility of African population structure and repeatedly caution against using patterns of genomewide sharing between modern and archaic humans is a strong hint that there are more things to come on the topic.

We should remember that the widely-circulated estimates of Neandertal->Eurasian introgression are based on genomewide averages. It is true that Reich et al. (2010) identified 13 regions of potential Neandertal introgression, which together make up a very small portion of the human genome. So, the jury is out on whether African population structure or Neandertal introgression is responsible for most of the genomewide pattern.

What you can be sure of is that many scientists are busy lining up full genomes from different human populations as we speak, and finding plenty of regions where haplotypes of extremely old divergence times co-exist in our species. We will probably learn more about such efforts during 2012.



Mol Biol Evol (2012)doi: 10.1093/molbev/msr301

Global genetic variation at OAS1 provides evidence of archaic admixture in Melanesian populations

Fernando L. Mendez, Joseph C. Watkins and Michael F. Hammer

Recent analysis of DNA extracted from two Eurasian forms of archaic human show that more genetic variants are shared with humans currently living in Eurasia than with anatomically modern humans in sub-Saharan Africa. While these genome-wide average measures of genetic similarity are consistent with the hypothesis of archaic admixture in Eurasia, analyses of individual loci exhibiting the signal of archaic introgression are needed to test alternative hypotheses and investigate the admixture process. Here, we provide a detailed sequence analysis of the innate immune gene, OAS1, a locus with a divergent Melanesian haplotype that is very similar to the Denisova sequence from the Altai region of Siberia. We re-sequenced a 7 kb region encompassing the OAS1 gene in 88 individuals from 6 Old World populations (San, Biaka, Mandenka, French Basque, Han Chinese, and Papua New Guineans) and discovered previously unknown and ancient genetic variation. The 5' region of this gene has unusual patterns of diversity, including 1) higher levels of nucleotide diversity in Papuans than in sub-Saharan Africans, 2) very deep ancestry with an estimated time to the most recent common ancestor of >3 million years, and 3) a basal branching pattern with Papuan individuals on either side of the rooted network. A global geographic survey of >1500 individuals showed that the divergent Papuan haplotype is nearly restricted to populations from eastern Indonesia and Melanesia. Polymorphic sites within this haplotype are shared with the draft Denisova genome over a span of ∼90 kb and are associated with an extended block of linkage disequilibrium, supporting the hypothesis that this haplotype introgressed from an archaic source that likely lived in Eurasia.

Link

Rapid onset of Aurignacian in Southwest France

Rapid onset of Aurignacian in Southwest France

Journal of Archaeological Science doi:10.1016/j.jas.2011.09.019

A Radiocarbon chronology for the complete Middle to Upper Palaeolithic transitional sequence of Les Cottés (France)

Sahra Talamo et al.

The Middle to Upper Palaeolithic transition is the key period for our understanding of Neanderthal and modern human interactions in Europe. The site of Les Cottés in south-west France is one of the rare sites with a complete and well defined sequence covering this transition period. We undertook an extensive radiocarbon dating program on mammal bone which allows us to propose a chronological framework of five distinct phases dating from the Mousterian to the Early Aurignacian at this site. We found that the Mousterian and Châtelperronian industries are separated from the overlying Protoaurignacian by a gap of approximately 1000 calendar years. Based on a comparison with Upper Paleolithic sites in Europe we see an overlap in the ages of Châtelperronian industries and Aurignacian lithic assemblages, which are usually associated with Anatomical Modern Humans, which is consistent with an acculturation at distance model for these late Neanderthals. The Proto and Early Aurignacian appear contemporaneous indicating that this transition was rapid in this region. Anatomically Modern Humans are present at the site of Les Cottés at least at 39,500 cal BP roughly coincident with the onset of the cold phase Heinrich 4.

Link

mtDNA and ethnic differentiation in East Africa

mtDNA and ethnic differentiation in East Africa

From the paper:

The pattern observed in East Africa (with the exception of the Khoisan-related Hadza and Sandawe populations), which combines a high level of within-population diversity with strong genetic structure among populations, suggests the occurrence of periodical episodes of admixture in these populations, separated by periods of isolation and genetic drift. Indeed, the observation of high levels of diversity within populations could be due to long-term large effective population sizes maintained in East Africa. In this case, however, little genetic structure between populations should be expected, since there would be little opportunity for genetic drift to act. Alternatively, gene flow can produce high within population diversity, and in the present case, it could also account for the extensive sharing of haplotypes and haplogroups observed between the Nyangatom and the Daasanach, as well as with other populations.

This seems like a very clever observation: substantial gene flow and a large effective population size would be inconsistent with population structure, as the different populations would be homogenized and drift would not be able to differentiate them. Long-term lack of gene flow, on the other hand, would not explain the sharing of haplotypes between populations, as each population would develop its own distinctive genetic signatures over time. Thus, the simplest explanation for the observed pattern is that gene flow has indeed occurred (accounting for the sharing of haplotypes), but that it was not continuous (accounting for the fact that populations are, after all, substantially differentiated).

From the paper:

The intermediate linkage disequilibrium (LD) found in East Africa (Tishkoff et al., 1996) in contrast with Europe (high LD) and Sub-Saharan Africa (low LD, Tishkoff & Kidd, 2004; Conrad et al., 2006), could be due to such admixture events, more frequently occurring in this region compared to other Sub-Saharan populations. Substantial levels of gene flow among Nilo-Saharan, Afro-Asiatic and Niger-Congo populations from Tanzania have already been inferred by Tishkoff et al. (2007a) and our results suggest that these gene flows could have occurred in a larger region extending up to Southern Ethiopia.

Indeed, in the absence of recent admixture, the East African populations would exhibit similar levels of LD with Sub-Saharan Africans., or even lower, as the indigenous East Africans are arguably older than those of the interior of the continent. The fact that they exhibit higher LD (intermediate between Europe and Sub-Saharan Africa) can be explained by admixture, i.e., the fact that they have inherited long stretches of DNA from the parental populations in each admixture event, and that time since that event has not been sufficiently long to cause the decay of these chunks into smaller pieces.

And, from the conclusions of the paper:

The high diversity in East Africa was interpreted as a sign of an ancient origin. However, our results might indicate that this high diversity could also come from a particular history of recent migrations and admixture promoted by the pastoralist societies that dominate in the region.

Note, that an East African origin of mankind is still the best hypothesis on palaeoanthropological and simply geographical grounds. However, the high genetic diversity found in East Africa does not necessarily reflect the antiquity of that population, but rather its history of repeated admixture by peoples of different origins.

There are two alternative hypotheses for why East Africans accumulated so much genetic diversity:

1. They are the oldest population, and have been accumulating genetic diversity for the longest period of time
2. They are substantially admixed with very divergent components (e.g., Semites, Nilo-Saharans, Cushitic speakers, and so on)

A not-so-bad example would be to compare them with other known population sources in the world, e.g., Anatolia, from where multiple waves of humans entered Europe in Paleolithic and Neolithic times. Many would agree that such movements took place, but it would be incorrect to see the population of Anatolia as a little-altered descendant of its earliest inhabitants, as the current genetic diversity observed there is -at least in part- the result of the settlement of the region by peoples from the Balkans, Central Asia, Levant, and even Western Europe.

Ann Hum Genet. 2009 Aug 25. [Epub ahead of print]

Genetic Evidence for Complexity in Ethnic Differentiation and History in East Africa.

Poloni ES, Naciri Y, Bucho R, Niba R, Kervaire B, Excoffier L, Langaney A, Sanchez-Mazas A.

Summary

The Afro-Asiatic and Nilo-Saharan language families come into contact in Western Ethiopia. Ethnic diversity is particularly high in the South, where the Nilo-Saharan Nyangatom and the Afro-Asiatic Daasanach dwell. Despite their linguistic differentiation, both populations rely on a similar agripastoralist mode of subsistence. Analysis of mitochondrial DNA extracted from Nyangatom and Daasanach archival sera revealed high levels of diversity, with most sequences belonging to the L haplogroups, the basal branches of the mitochondrial phylogeny. However, in sharp contrast with other Ethiopian populations, only 5% of the Nyangatom and Daasanach sequences belong to haplogroups M and N. The Nyangatom and Daasanach were found to be significantly differentiated, while each of them displays close affinities with some Tanzanian populations. The strong genetic structure found over East Africa was neither associated with geography nor with language, a result confirmed by the analysis of 6711 HVS-I sequences of 136 populations mainly from Africa. Processes of migration, language shift and group absorption are documented by linguists and ethnographers for the Nyangatom and Daasanach, thus pointing to the probably transient and plastic nature of these ethnic groups. These processes, associated with periods of isolation, could explain the high diversity and strong genetic structure found in East Africa.

Link