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Category Archives: Namibia

SW African Bantu matrilineages

Prolific researcher Chiara Barbieri has put online another interesting study on African genetics, this time about the Bantu populations of Southwestern and Central-Southern Africa (i.e. Namibia, Angola, Botswana and Zambia).
Chiara Barbieri et al., Migration and interaction in a contact zone: mtDNA variation among Bantu-speakers in southern Africa. bioRXiv 2014. Freely accessible (pre-pub) → LINK

ABSTRACT

Bantu speech communities expanded over large parts of sub-Saharan Africa within the last 4000-5000 years, reaching different parts of southern Africa 1200-2000 years ago. The Bantu languages subdivide in several major branches, with languages belonging to the Eastern and Western Bantu branches spreading over large parts of Central, Eastern, and Southern Africa. There is still debate whether this linguistic divide is correlated with a genetic distinction between Eastern and Western Bantu speakers. During their expansion, Bantu speakers would have come into contact with diverse local populations, such as the Khoisan hunter-gatherers and pastoralists of southern Africa, with whom they may have intermarried. In this study, we analyze complete mtDNA genome sequences from over 900 Bantu-speaking individuals from Angola, Zambia, Namibia and Botswana to investigate the demographic processes at play during the last stages of the Bantu expansion. Our results show that most of these Bantu-speaking populations are genetically very homogenous, with no genetic division between speakers of Eastern and Western Bantu languages. Most of the mtDNA diversity in our dataset is due to different degrees of admixture with autochthonous populations. Only the pastoralist Himba and Herero stand out due to high frequencies of particular L3f and L3d lineages; the latter are also found in the neighboring Damara, who speak a Khoisan language and were foragers and small-stock herders. In contrast, the close cultural and linguistic relatives of the Herero and Himba, the Kuvale, are genetically similar to other Bantu-speakers. Nevertheless, as demonstrated by resampling tests, the genetic divergence of Herero, Himba, and Kuvale is compatible with a common shared ancestry with high levels of drift and differential female admixture with local pre-Bantu populations.

Figure 1: Map showing the rough geographical location of populations, 
colored by linguistic affiliation. Abbreviations of population labels are 
as specified in Table 1.

In spite of the Bantu-centric approach of the study, which also has its merits, my greatest interest is rather in the less typically Bantu lineages, which speak of admixture with several pre-Bantu populations.
In this sense I find the following highlights:

Fig. S2 (annotated in green by Maju): CA plots based on haplogroup frequencies. Left: all the dataset, right: excluding outliers.

L3d and L3f founder effect:
The Himba and Herero, as well as the non-Bantu pastoralists Damara make one distinctive cluster defined by the high frequencies of haplogroup L3d, as well as L3f (not present among the Damara but found among the Kuvale). As discussed in the paper, the Himba and Herero may be related to the Kuvale of SW Angola but they have notable differential levels (or directionality) of aboriginal admixture. 
As both L3d and L3f are present in West and East Africa alike, it is interesting to track the specific subhaplogroups implicated in this founder effect, something done in fig. 4. 
The main L3d sublineage is L3d3a1, whose haplotype network shows a largely Khoisan centrality (not Damara) although this node is shared also by some unspecified “other Bantu”. The Southern Africa specificity of L3d3a was already noticed in the past (see here). So it is very possible that we are before an aboriginal Southern African lineage, maybe arrived with the first Khoisan Neolithic (or whatever other ancient flow) rather than a Bantu-specific founder effect. 
The main L3f subhaplogroup is L3f1b4a, which seems more specifically Bantu, with a major branch concentrated among the Himba, Herero and Kuvale. This lineage is not found among the Damara in spite of the other strong affinity of this Khoisan population towards the Himba and Herero. L3f1b is found in Southern Africa, Kenya and Oman (per Bihar 2008), so we are probably before a distinctive East African element, not too likely to be genuinely Bantu but possibly just assimilated into Bantu ethnic identity. 
Even if both lineages converge in the Himba and Herero, they are almost certainly different inputs, one of Damara (herder Khoisan) origin and the other of Bantuized East African origin maybe.
L1b founder effect:
L1b is essentially a West African lineage concentrated in the Sahel area from Chad westwards (although L1b1a2 is from the Nile basin). A particularly high frequency population are the Fulani pastoralists, original from the Westernmost African plateaus, who ruled many kingdoms in West Africa between the collapse of the colonial rule by Morocco and the consolidation of the European conquest of the continent.
As this study does not dwell in sublineages, we cannot understand the most likely specific origins of it among several Southern African populations, specifically the pooled NE Zambians (13%) and the Fwe and Shanjo of SW Zambia (24-27%).
In any case it is a notorious founder effect, almost absent in other Bantus of the area (0-10%).
Typical L0d Khoisan admixture:
This element is concentrated in Botswana (~25%) and with highest frequencies in the SW Kgalagadi (53%). It is also important among the Kuvale of SW Angola (21%). Other Bantu populations in this dataset have frequencies under 10%, some even zero. The Damara have 13%.
We know from previous studies that it is also found at high frequencies among the Xosha of South Africa (L0d3).
While L3h appears marked in the graph, the lineage is in fact absent in all populations except at very low frequency among the Kuvale (2%), so it does not seem actually of any relevance. 
Less typical L0k around SW Zambia:
While L0k is generally considered an aboriginal Southern African lineage it has a much more northernly distribution than the more common and surely older L0d. Its area of greatest commonality seems to be SW Zambia (see here and here).
This study confirms this distribution:

Supplementary Figure S3[A]: Haplogroup frequencies of important haplogroups in the populations studied here. A: Haplogroups L0d and L0k.(…)

The size of the circles is proportional to the sample size.

High frequencies of L1c (Pygmy admixture marker) among Southern African Bantus:
An interesting element is the commonality of L1c, typical of Western Pygmies and some other populations from Gabon (possibly representative of the wider West-Central Africa jungle region, not too well studied otherwise), among almost all Bantu populations in this dataset. 
The exceptions are the Herero, Himba, Kgalagadi and Tswana (0%), as well as the NE Zambians (4%). All the rest have frequencies between 12% and 30%. Even the non-Bantu Damaras have 11% of it.
In my understanding this almost certainly implies a notable level of admixture with Western Pygmies of the Bantus from especially Angola and West Zambia. A phenomenon that may be widespread in Central-West Africa. 
It is notable however that at least many of the populations with the highest likely Khoisan admixture (in its various forms, discussed in the previous sections) have the lesser frequencies of L1c (Pygmy admixture). So to a great extent these two aboriginal influences in Bantu mtDNA seem mutually exclusive and were probably produced after settlement rather than “on the march”. 
This in turn arises some interesting questions about the ethnic geography of Africa before the Bantu expansion. 

Update: I just noticed that Ethiohelix has parsed the haplogroups’ frequency into a very helpful chartLINK.

See also:
 

Khoesan and Coloured autosomal DNA in context

There has been a number of studies coming out recently on Khoesan genetics but this one does not seem to be just redundant, providing some extra information instead.

Desiree C. Petersen et al., Complex Patterns of Genomic Admixture within Southern Africa. PLoS Genetics 2013. Open accessLINK [doi:10.1371/journal.pgen.1003309]


Abstract


Within-population genetic diversity is greatest within Africa, while between-population genetic diversity is directly proportional to geographic distance. The most divergent contemporary human populations include the click-speaking forager peoples of southern Africa, broadly defined as Khoesan. Both intra- (Bantu expansion) and inter-continental migration (European-driven colonization) have resulted in complex patterns of admixture between ancient geographically isolated Khoesan and more recently diverged populations. Using gender-specific analysis and almost 1 million autosomal markers, we determine the significance of estimated ancestral contributions that have shaped five contemporary southern African populations in a cohort of 103 individuals. Limited by lack of available data for homogenous Khoesan representation, we identify the Ju/’hoan (n = 19) as a distinct early diverging human lineage with little to no significant non-Khoesan contribution. In contrast to the Ju/’hoan, we identify ancient signatures of Khoesan and Bantu unions resulting in significant Khoesan- and Bantu-derived contributions to the Southern Bantu amaXhosa (n = 15) and Khoesan !Xun (n = 14), respectively. Our data further suggests that contemporary !Xun represent distinct Khoesan prehistories. Khoesan assimilation with European settlement at the most southern tip of Africa resulted in significant ancestral Khoesan contributions to the Coloured (n = 25) and Baster (n = 30) populations. The latter populations were further impacted by 170 years of East Indian slave trade and intra-continental migrations resulting in a complex pattern of genetic variation (admixture). The populations of southern Africa provide a unique opportunity to investigate the genomic variability from some of the oldest human lineages to the implications of complex admixture patterns including ancient and recently diverged human lineages.

The array of Khoesan populations senso stricto analyzed in this study is much smaller than that of Schebusch 2010 but this study has the advantage of including Cape Coloureds and their Baster relatives, partially descendants from the otherwise extinct pastoralist Khoekhoe (Hottentots, now considered a derogative term) who lived in much of Southern Africa upon the arrival of Bantu and Europeans, as well as the amaXhosa, a Bantu people which clearly display marked Khoesan admixture.

Figure 1. Map of southern Africa
showing distribution of sampling per population identifier and
significant historical events that likely shaped ancestral
contributions.

There is brief mention of maternal and paternal DNA. Just to mention that mtDNA being mostly aboriginal (L0d/L0k) among the Khoesan (86-100%), the Coloureds (68%) and even the Xhosa (47%, all L0d), while aboriginal Y-DNA (essentially A2b and A2c2, plus occasional B2) is concentrated among the Ju/’hoan, with the !Xun being instead dominated by E1b1-M275, of putative East African (Nilotic?) origins. This is consistent with the !Xun being historically pastoralists. European patrilineages, notably R1b, are dominant among the Baster (92%) and Cape Coloured (71%).
Coloureds only make up some 9% of South African population but they dominate the countryside in much of the former Cape Province. Namibian Basters are a subset of them who migrated northwards in 1868.

Figure 2.  PCA and STRUCTURE analysis (click to expand)
We can see in the graphics above how the North Cape Coloured and Baster only display minor Bantu admixture, being essentially a variable mix of European and Khoesan ancestry, with probably also some Malay input (apparent in the increase of the blue component relative to the European reference). Instead East Cape and Cape Town (D6) Coloured appear to have greater apportion of Bantu ancestry and, especially the later, a notable increase of the East Asian input.
The STRUCTURE graph, particularly at K=9, is also informative about other African populations but I won’t dwell in that here. 
The authors also made an interesting exercise of analysis using Ancestry Informative Markers with the !Xun and Xhosa:

Figure 4. Ju/’hoan-Yoruba ancestry
informative markers (AIMs) defined ancestral contributions to the !Xun
and amaXhosa, providing evidence for two distinct !Xun lineages with
differing ancestral contributions.
It seems evident that much of the !Xun ancestry (up to 70%) does not fall in either (Ju/’hoan-Yoruba) category but it is something else, probably specific to this people. The Xhosa Khoesan ancestry also seems closer to the pastoralist !Xun than to the (likely more genuinely ancient) Ju/’hoan. 

There is some more info in the paper but I feel that the essentials are sufficiently covered here. 

See also:
 

Khoe-San matrilineages and prehistory

A most interesting study has just been published that reconstructs the prehistory of the Khoe-San peoples of Southern Africa primarily using mitochondrial DNA analysis but with very important reliance on archaeological data as well.
Karina M. Schlabusch et al., MtDNA control region variation affirms diversity and deep sub-structure in populations from Southern Africa. BMC Evolutionary Biology 2013. Open accessLINK [doi:10.1186/1471-2148-13-56]

Abstract (provisional)


Background

The current San and Khoe populations are remnant groups of a much larger and widely dispersed population of hunter-gatherers and pastoralists, who had exclusive occupation of southern Africa before the influx of Bantu-speakers from 2 ka (ka = kilo annum [thousand years] old/ago) and sea-borne immigrants within the last 350 years. Here we use mitochondrial DNA (mtDNA) to examine the population structure of various San and Khoe groups, including seven different Khoe-San groups (Ju/’hoansi, !Xun, /Gui+//Gana, Khwe, =Khomani, Nama and Karretjie People), three different Coloured groups and seven other comparative groups. MtDNA hyper variable segments I and II (HVS I and HVS II) together with selected mtDNA coding region SNPs were used to assign 538 individuals to 18 haplogroups encompassing 245 unique haplotypes. Data were further analyzed to assess haplogroup histories and the genetic affinities of the various San, Khoe and Coloured populations. Where possible, we tentatively contextualize the genetic trends through time against key trends known from the archaeological record.

Results

The most striking observation from this study was the high frequencies of the oldest mtDNA haplogroups (L0d and L0k) that can be traced back in time to ~100 ka, found at high frequencies in Khoe-San and sampled Coloured groups. Furthermore, the L0d/k sub-haplogroups were differentially distributed in the different Khoe-San and Coloured groups and had different signals of expansion, which suggested different associated demographic histories. When populations were compared to each other, San groups from the northern parts of southern Africa (Ju speaking: !Xun, Ju/’hoansi and Khoe-speaking: /Gui+//Gana) grouped together and southern groups (historically Tuu speaking: =Khomani and Karretjie People and some Coloured groups) grouped together. The Khoe group (Nama) clustered with the southern Khoe-San and Coloured groups. The Khwe mtDNA profile was very different from other Khoe-San groups with high proportions of Bantu-speaking admixture but also unique distributions of other mtDNA lineages.

Conclusions

On the whole, the research reported here presented new insights into the multifaceted demographic history that shaped the existing genetic landscape of the Khoe-San and Coloured populations of southern Africa.

From the reading of the paper, I gather the following chronology (which should be always taken with some caution because of the uncertainty of “molecular clock” methods but in this case they seem reasonably backed from the material/cultural evidence record):
  1. L0d coalescence time estimate may correlate with the arrival of MSA to the region c. 100 Ka ago (I estimated once ~90 Ka, so it is consistent with my thought).
  2. Its sublineage L0d1’2 might have expanded c. 50 Ka ago (I would rather think of a more ancient chronology, soon after the L0d node – they can’t correlate it properly with any obvious archaeological pattern, so it might be, I guess, more related to the apogee of MSA c. 75 Ka ago).
  3. Some L0d1 subclades (notably L0d1a, L0d1b) would have expanded with the transition to LSA (40-20 Ka ago).
  4. L0d2a shows an star-like expansion that they estimate to have happened c. 7-8 Ka ago and would be related to an Epipaleolithic (with microlithic industry) that is also notable for the increase of the density of archaeological findings in South Africa and Lesotho. This lineage also shows secondary expansion with pastoralism later on.
  5. The introduction of herding c. 2000 years ago may have affected the correlations between the various L0d lineages. However most lineages show signs of expansion in this period. The main exception is L0d1a (decrease instead) and to some extent L0d1c (first decrease, later increase probably related to the !Xun late adoption of pastoralism, affecting especially to L0d1c1).
    1. L0d3 is too old to have expanded with pastoralism, so the authors reject  Tatiana Karafet’s hypothesis that it expanded in this period and that it could be related (to most unlikely) linguistic relation between Sandawe and Khoe-San. Instead they suggest (as I did in the past) that L0d3 had an East African distribution instead with only minor spreading to the Khoe-San in relation with pastoralism.
  6. The recent Iron Age (last millennium) arrival of Bantu-speakers absorbed primarily L0d2a, which is the most common lineage of Khoe-San peoples (including Coloureds, with the partial exception of Cape Coloured, where it is second to L0d2b).
The paper only briefly mentions L0k1, which is most concentrated towards Katanga (D.R. Congo) and may therefore have arrived to Southern Africa only with Bantu or pastoralist flows.
Frequencies and estimated timelines of major Southern African L0d and L0k lineages (from fig. 4):

See also:

 

The Neolithic site of Leopard Cave (Namibia)

A new interesting open access paper on the Neolithic of Southern Africa has been published:

Abstract
The origins of herding practices in southern Africa remain controversial. The first appearance of domesticated caprines in the subcontinent is thought to be c. 2000 years BP; however, the origin of this cultural development is still widely debated. Recent genetic analyses support the long-standing hypothesis of herder migration from the north, while other researchers have argued for a cultural diffusion hypothesis where the spread of herding practices took place without necessarily implicating simultaneous and large population movements. Here we document the Later Stone Age (LSA) site of Leopard Cave (Erongo, Namibia), which contains confirmed caprine remains, from which we infer that domesticates were present in the southern African region as early as the end of the first millennium BC. These remains predate the first evidence of domesticates previously recorded for the subcontinent. This discovery sheds new light on the emergence of herding practices in southern Africa, and also on the possible southward routes used by caprines along the western Atlantic coast.

The caprine (sheep) remains are the oldest ones of the whole region, being dated to c. 2270 BP, almost 200 years (est.) before the next oldest one with a good date (Spoegrivier, Western South Africa, dated to c. 2100 BP). Notice however the “uncertain” much older date for the Orowanjo site near the Angolan border (c. 3100 BP) in the map below:

Fig. 4Later Stone Age sites of southern Africa with early evidence of caprines
Caprines are a focus of this research but they are only a minority (2) of the animals identified in Leopard Cave, with bovines (cows) being the most important (16) among domesticates and in general if we exclude ostrich eggshells. Unspecified rodents and birds reach similar figures to those of bovine cattle but these belong to diverse species with all likelihood. 
Besides animal remains the cave has also yielded a dearth of stone artifacts and some pottery. 
It is unclear to which modern ethnic group, if any, can the site be associated with but the area has been traditionally inhabited by the Damara people, who speak either Khoekhoe or sometimes Herero languages but who, like their peculiar Northern Neighbors, the Himba people, are generally speculated to be a different ethnic, cultural and probably also genetic stock from both Khoisan and Bantu peoples. Sadly I do not know of any genetic study on either ethnic group, which may be of great interest for ethnographic and prehistoric reconstruction purposes.

Update (Jul 29): for a new study on Khoesan (and also Damara, Himba, etc.) autosomal genetics see HERE.

 
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Posted by on July 12, 2012 in Africa, Namibia, Neolithic, Prehistory