The Origin of Paternal Haplogroups C, F, D, & E
Origin of Modern humans
Jebel Irhoud 1 (skull of a basal modern human/ basal Homo sapiens) and its reconstruction by Kennis & Kennis [Link]
The oldest Homo sapiens fossils were found in Jebel Irhoud - Morocco, which is in North Africa; they are estimated to be approximately 300,000 years old. They have an intermediate set of traits between those of Homo heidelbergensis - from which Homo sapiens evolved - and those of fully modern Homo sapiens: their faces project forward less than those of Homo heidelbergensis but more than those of fully modern Homo sapiens; they have very large brow ridges like Homo heidelbergensis, which is unlike fully modern Homo sapiens; and their craniums lack the occipital bun of Homo heidelbergensis but also lack the globular shape that characterizes fully modern Homo sapiens. These fossils’ intermediate morphology indicates that they were at the beginning stages of our species’ birth; therefore, they are referred to as basal Homo Sapiens or basal modern humans.
Calvaria of Omo I (left) and that of Omo II (right)
The second oldest Homo sapiens fossils were found in Ethiopia, which is in East Africa; they include Omo I and Omo II and have been estimated to be 233,000 years old. Apart from a few archaic traits, such as minimally pronounced brow ridges and minimal midfacial projection, they have practically all of the traits of fully modern humans; they possess our species’ characteristic globular brain case, defined chin, and vertical forehead. Therefore, they are referred to as anatomically modern humans.
Skhul V and Qafzeh 9 and reconstructions of them by Kennis & Kennis [Link]
The third oldest Homo sapiens fossils were found in Israel, which is just outside of Northeast Africa; they include Skhul V and Qafzeh 9, and they have been estimated to be 80,000 to 120,000 years old. They are referred to as anatomically modern humans as well.
So, it seems that Homo sapiens originated in North Africa and then some members of the species spread to East Africa, where they evolved into anatomically modern humans (while others stayed in North Africa and became a different subspecies of Homo sapiens: Aterians [Link]). Afterwards, anatomically modern humans started migrating out of Africa via the Northeast region of the continent (i.e. Egypt).
The evolutionary development of anatomically modern humans in East Africa is bolstered by the discovery of additional but taxonomically different fossils in Ethiopia: those of Herto Man, otherwise known as Homo sapiens idaltu, which are estimated to be 160,000 years old.
Skull of Herto Man / Homo sapiens idaltu (BOU-VP-16)
That this other subspecies of Homo sapiens has been found in Ethiopia indicates the following: the ancestral population that produced both anatomically modern humans and Homo sapiens idaltu settled in East Africa and remained there for a couple hundred thousand years - long enough to diverge into two subspecies. This is far more likely than both subspecies evolving in geographically separate regions and then coincidentally traveling to the same general area in such a large continent.
Origin of Major modern human paternal haplogroups
Based on the aforementioned evidence, it is apparent that all modern populations ultimately originated in East Africa. This raises the question of whether paternal haplogroups that exist exclusively outside of Africa or that exist in both Africa and outside of it formed and coalesced into their most recent common ancestors (MRCAs) before or after the ancestors of non-Africans left East Africa. This can be determined by analyzing the dates at which they formed and the dates of their MRCAs, and by comparing these dates to the estimated time period during which the ancestors of non-Africans migrated out of Africa.
According to YFull:
NOTE: YBP means Years Before Present.
Generation 1: 88,000 to 68,500 YBP
CT: formed 88,000 YBP | MRCA 68,500 YBP
Generation 2: 68,500 to 65,200 YBP
CF: formed 68,500 YBP | MRCA 65,900 YBP
DE: formed 68,500 YBP | MRCA 65,200 YBP
Generation 3: 65,900 to 46,500 YBP
C (C-M130): formed 65,900 YBP | MRCA 48,800 YBP
F (F-M89): formed 65,900 YBP | MRCA 48,800 YBP
D (D-CTS3946): formed 65,200 YBP | MRCA 46,500 YBP
E (E-M96): formed 65,200 YBP | MRCA 52,300 YBP
NOTE:
CT -> CF & DE
CF -> C & F
DE -> D & E
The migration of anatomically modern humans out of Africa that produced non-Africans is estimated to have occurred 50,000 YBP to 70,000 YBP, as stated in numerous peer-reviewed scientific articles, such as the following two.
A Rare Deep-Rooting D-Zero African Y-Chromosomal Haplogroup and Its Implications for the Expansion of Modern Humans Out of Africa (2019) [Link]: Present-day humans outside Africa descend mainly from a single expansion out ∼50,000–70,000 years ago, but many details of this expansion remain unclear, including the history of the male-specific Y chromosome at this time.
A recent bottleneck of Y chromosome diversity coincides with a global change in culture (2015) [Link]: The estimated time line of the Y chromosome coalescent events in non-African populations (Supplemental Fig. S9) fits well with archaeological evidence for the dates of colonization of Eurasia and Australia by anatomically modern humans as a single wave ∼50 kya.
Therefore, CT's formation can be confidently placed in Africa because it formed 88,000 YBP, which was before the most distant time at which the migration is estimated to have occurred (70,000 YBP).
However, if 70,000 YBP was indeed when the migration occurred, then CF, DE, C, F, D, and E all must have formed outside of Africa, because they all formed after 70,000 YBP. Conversely, if the closer time period of 50,000 YBP was when the migration occurred, then they all must have formed inside of Africa, because they all formed before 50,000 YBP. Complicating matters further is the range of time between 50,000 YBP and 70,000 YBP, because the migration could have occurred at any point therein as well.
However, evolutionary geneticists William Amos and Joe Hoffman of the University of Cambridge conducted an analysis of genomes from 53 modern human populations, thereby determining a narrower period of time during which the migration could have occurred. Particular characteristics of these genomes indicated that the ancestors of non-Africans experienced a population bottleneck 50,000 to 60,000 years ago as they migrated out of Africa.
This is explained in the following quote from the article How We Lost Our Diversity [Link]:
With the recent publication of a large data set of 763 microsatellite markers--short stretches of DNA that are repeated in the genome--from 53 populations in the Human Genome Diversity Project, evolutionary geneticists William Amos and Joe Hoffman of the University of Cambridge in the United Kingdom had enough genomic data to test both models. Using a software program called BOTTLENECK, the two searched for the signature of bottlenecks or founder effects in each population to see if the loss of diversity occurred suddenly or gradually as adjacent populations moved farther from Africa. Different genetic markers of diversity decline at different rates, creating an imbalance…. The team uncovered strong signs of this imbalance between rare alleles and heterozygosity in two populations--one, in people living today in the Middle East, and the other in the Yakut, who live near the Bering Strait. That indicated to the researchers that the first bottleneck occurred as people migrated out of Africa to the Middle East about 50,000 to 60,000 years ago…. What caused the bottlenecks? Amos suggests that obstacles--such as the sea, glaciers, or mountains--on the route from Africa to the Middle East, across the Himalayas and over the Bering Strait, held many migrants back, so that a much smaller group moved on, producing offspring to inhabit new parts of the globe.
The findings of the geneticists mentioned in the above quote cut the previously mentioned time period during which the migration could have occurred in half; they reduce 50,000 to 70,000 years ago down to 50,000 to 60,000 years ago. However, this is still a wide range.
To pinpoint the date at which the migration occurred, the most recent common ancestor of each paternal haplogroup must be considered. By comparing them, a trend becomes apparent: the MRCAs of C, F, and D are the only MRCAs who lived after the most recent time at which the migration is estimated to have occurred (50,000 YBP), while the MRCAs of all of the other haplogroups lived before this time - particularly the MRCA of Haplogroup E, who lived 52,300 YBP. This is consistent with the current distribution of paternal haplogroups worldwide: subclades of C, F, and D are present in Eurasia, Oceania, and the Americas but are absent from Africa (apart from D0 [D Zero], which is a very early branch of D that has been discovered in less than a handful of Africans); however, subclades of E are present in both Africa and Eurasia, but they are overwhelmingly so in Africa and barely so in Eurasia - apart from subclades of E1b1b (E-M215), which shall be discussed later herein. Therefore, the date of the migration was likely 50,000 YBP.
The following list of events are based on the aforementioned dates at which the descendant haplogroups of CT formed and the dates of their MRCAs, as well as the logically deduced date of the migration out of Africa:
CF formed C & F inside of Africa 65,900 YBP.
DE formed D & E inside of Africa 65,200 YBP.
There was a population (perhaps a federation of tribes) that included all males who possessed C, F, D, and E: Population CFDE
Population CFDE split into two populations:
Population CFD, which contained all of the males who possessed subclades of C, F, and D.
Population E, which contained all of the males who possessed subclades of E.
Population CFD migrated out of Africa 50,000 YBP.
Population E remained in Africa when Population CFD migrated out of Africa.
Sometime after 46,500 YBP (the time at which the MRCA of D lived), males possessing D0 [D Zer0] migrated “back” to Africa. Due to the date of D’s MRCA being 46,500 YBP, he must have lived outside of Africa since the migration out of Africa occurred 50,000 YBP; therefore, the presence of D0 [D Zero] in Africa must be due to a migration of D0 [D Zero] back to Africa.
One might be inclined to think that if D0’s [D Zero’s] presence in Africa can be explained via a migration from Eurasia back to Africa, then so can the presence of all of the subclades of E that currently exist in the continent. However, the different frequencies and diversities of these two sets of haplogroups therein make a return from Eurasia likely for the former but unlikely for the latter.
D0 [D Zero] is the only subclade of D that has been discovered in Africa, and it has been found in only three African men (from Nigeria) [source]; therefore, it is extremely rare and extremely homogenous in the continent. The extremely low frequency and extreme homogeneity of D therein strongly imply that the subset of D-carrying males who either remained or returned was indeed small. Additionally, because the MRCA of D lived after the migration, this small subset of males were not born in the continent and therefore “returned” to it.
Conversely, E occurs most frequently and is most diverse by far in Africa. The males who possess its subclades therein amount to hundreds of millions, and its oldest subclades to its youngest ones can be found therein. However, as already stated, only one of its subclades occurs frequently outside of the continent: E1b1b. Additionally, E1b1b descends from subclades that are rare outside but abundant inside of the continent, and the regions outside of the continent in which it’s strongly present are nearby (i.e. the Mediterranean Basin and the Arabian Peninsula); this implies radiation from Africa. Consider the following quote from a peer-reviewed scientific article.
Saudi Arabian Y-Chromosome diversity and its relationship with nearby regions [Link]: Around 14% of the Saudi Arabia Y-chromosome pool is typical of African biogeographic ancestry … Global male inputs from Sub-Saharan Africa and Asia across Iran, not the Levant, into the Arabian Peninsula have been estimated in this study, as 13.4% and 16.6% from both source areas respectively…. Without dismissing the role mediated by slavery, the geographical distribution of these sub-Saharan African lineages in the Arabian Peninsula seems to indicate a prehistoric entrance of a noticeable portion of these lineages that participated in the building of the primitive Arabian population…. The data confirm that this area has mainly been a recipient of gene flow from its African and Asian surrounding areas, probably mainly since the last Glacial maximum onwards.
Furthermore, E1b1b formed 41,400 YBP, which was 10,900 years after the MRCA of E lived (52,300 YBP). Therefore, if E had been in Eurasia from at least the time of its MRCA to the time at which E1b1b formed, then it would have been populating Eurasia and diversifying therein for 10,900 years!
Alternatively, if there still existed other lineages of E aside from those of E’s MRCA when E1b1b formed, then E would have possibly been in Eurasia since its own formation 65,200 YBP; this means that it would have been populating Eurasia and diversifying therein possibly for as long as 23,800 years!
In either case, it would have grown to a population of tens-or-hundreds of thousands of males and diversified to dozens-or-hundreds of subclades.
Therefore, a migration from Eurasia to Africa of E would have required tens-or-hundreds of thousands of males (and their female companions) to collectively decide to migrate to Africa. As this migration would have occurred thousands or tens-of-thousands of years ago, when everyone lived as hunter-gatherers who were divided into bands of approximately 50 people at most - and which didn’t engage in complex inter-group coordination - the odds of this having happened are extremely low. Additionally, even if such a migration were the result of people fleeing from a natural disaster or escaping food scarcity, the odds of practically all carriers of E migrating to the same region are very low - especially when considering that even the carriers of any one particular haplogroup out of C, F, and D migrated sporadically relative to fellow carriers of the same haplogroup. For example, subclades of Paternal Haplogroup C can be found in Australia, East Asia, and the Americas (among Native Americans); and haplogroups that descend from F can be found in the Near East, Europe, and Asia.
As a side note, it must be considered that all major migrations of haplogroups in pre-history must have been the result of uncoordinated movements of populations in response to natural disasters, food scarcity, and drifting migratory routes of prey animals. Subsequently, the distribution of haplogroups was somewhat random; this explains why D is found in Africa and as distantly as East Asia - or why C, as already stated, is found in distantly separated locations such as Australia, East Asia, and the Americas (among Native Americans).
As an additional side note, it must also be considered that a migration by the Out of Africa population (Population CFD) from Africa to Eurasia 50,000 YBP would not have been impeded by the population being too large, since the population suffered a bottleneck event, as explained in an article cited previously.
E-M215 / E1b1b: the anomalous subclade of E-m96
E1b1b is an anomalous subclade of E, because it is the only subclade of E that is abundantly present outside of Africa, specifically in Eurasia. However, like all other subclades of E, it originated in Africa, as shall be proven herein.
The oldest samples of E1b1b have been found in remains of the ancient Iberomaurusians of Taforalt - Morocco, as well as in remains of the ancient Natufians of the Near East. Both of these populations’ ancestries were predominantly Eurasian, but they both had sizeable percentages of indigenous African ancestry (as shall be explained). Additionally, the Iberomaurusians of Taforalt had maternal haplogroups that are innately associated with Eurasians and which they therefore must have inherited from their Eurasian ancestors; however, their paternal haplogroups, which are subclades of E1b1b could have come from their indigenous African ancestors because they are not innately associated with Eurasians. This conclusion can be pieced together from the following pieces of information.
The Iberomaurusians of Taforault
A Reconstruction of an Iberomaurusian man by Ancestral Whispers [Link].
The Taforalt specimens are estimated to have lived between 15,100 YBP and 13,900 YBP, as stated in the following quote from the peer-reviewed article Pleistocene North African genomes link Near Eastern and sub-Saharan African human populations (2018) [Link]: Here we present genome-wide data from seven individuals, directly dated between 15,100 and 13,900 calibrated years before present (cal. yr B.P.), from Grotte des Pigeons near Taforalt in eastern Morocco.
Because they themselves lived in ancient times, the source of their paternal haplogroup also lived in ancient times.
The autosomal DNA of the Taforalt specimens whose genomes were analyzed was determined to be 63.5% Eurasian and 36.5% Sub-Saharan African, as stated in another quote from the aforementioned article: A two-way admixture model, comprising Natufian and sub-Saharan African populations, does not significantly deviate from our data, with 63.5% Natufian and 36.5% sub-Saharan African ancestry.... West Africans, such as Mende and Yoruba most strongly pull out the sub-Saharan African ancestry in Taforalt.
The Natufians were a population who inhabited the Levant (Middle East) as early as 14,000 YBP (12,000 BC). Even though they possessed 6% indigenous African ancestry (as shall be explained), they can be used to model the Eurasian ancestry of the Taforalt specimens due to being 94% Eurasian.
The maternal haplogroups of the Taforalt specimens whose genomes were analyzed are U6a and M1b, and they are both likely of Eurasian origin, as stated in another quote from the aforementioned article: Mitochondrial consensus sequences of the Taforalt individuals belong to the U6a (six individuals) and M1b (one individual) haplogroups, which are mostly confined to present-day populations in North and East Africa. U6 and M1 have been proposed as markers for autochthonous Maghreb ancestry, which might have been originally introduced into this region by a back-to-Africa migration from West Asia.
All maternal haplogroups that are not labeled as an “L” haplogroup are Eurasian; they are labeled with letters other than “L” to signify that they are associated with the Out of Africa population (though they ultimately descend from Africa’s Maternal Haplogroup L3). Therefore, when the quote says that U6 and M1 could possibly be markers for autochthonous (native) Maghreb ancestry (the Maghreb is Northwest Africa), what it means is that they could have formed in Northwest Africa even though they ultimately descend from maternal haplogroups that formed in Eurasia; conversely, when the quote says that Maghreb ancestry - with which the two haplogroups are associated - might be from West Asia, it means that the two haplogroups may have formed in West Asia and then were brought to Northwest Africa later. In short, whether these two haplogroups formed in West Asia or formed in North Africa, they are Eurasian in origin; their lineages descend from people who participated in the Out of Africa migrations and whose descendants evolved to adapt to Eurasia.
As the Iberomaurusians had Eurasian ancestry, it is unquestionable that their Eurasian ancestors were the source of their Eurasian maternal haplogroups.
The paternal haplogroup of the Taforalt specimens whose genomes were analyzed is E-M78 / E1b1b1a1, as stated in yet another quote from the aforementioned article: Consistently, we find that all males with sufficient nuclear DNA preservation carry Y haplogroup E1b1b1a1 (M-78).
As the Iberomaurusians’ paternal haplogroup is not innately associated with the Out of Africa population, it is logical to conclude that their indigenous African ancestors may have been the source of their paternal haplogroup, which is a subclade of E1b1b. Considering this and considering the predominance of indigenous Africans in regard to the overwhelming majority of Haplogroup E’s other subclades, it is logical to conclude that E1b1b originated among indigenous Africans.
The Natufians
A Reconstruction of a Natufian man by Ancestral Whispers [Link].
As stated above, the Natufians inhabited the Levant as early as 14,000 YBP (12,000 BC); this is stated in the peer-reviewed scientific article Genomic insights into the origin of farming in the ancient Near East (2016) [Link]: We report genome-wide ancient DNA from 44 ancient Near Easterners ranging in time between ~12,000 [bc] and 1,400 bc, from Natufian hunter-gatherers to Bronze Age farmers.
The paternal haplogroup of the Natufians whose genomes have been analyzed is E1b1b1b2-Z830, which is stated in the peer-reviewed scientific article Placing Ancient DNA Sequences into Reference Phylogenies (2022) [Link]: Apart from those described above, the majority (n = 11) of other East African PN samples were placed in E1b1b1b2b-V1515 lineages, a sub lineage of E1b1b1b2-Z830 found in the Levantine proto-agriculturalist Natufians…
The autosomal DNA of Natufian specimens whose genomes were analyzed was determined to be a mixture of several Eurasian sources and one indigenous African source (Omotic, which is indigenous to East Africa). Their Omotic ancestry was a small component of their genome, but it was larger before the influx of Arabian ancestry into their genome during the Neolithic, according to the scientific article Re-analysis of Whole Genome Sequence Data From 279 Ancient Eurasians Reveals Substantial Ancestral Heterogeneity [Link]: The Natufian sample consisted of 61.2% Arabian, 21.2% Northern African, 10.9% Western Asian, and 6.8% Omotic ancestry. The transition in the Levant from the Epipaleolithic to the Neolithic period involved an increase of Arabian ancestry at the expense of Northern African and Omotic ancestries.
The same article posits that the paternal haplogroup of the Natufians (E1b1b1b2-Z830) and Haplogroup E as a whole may have come from their Omotic ancestry: Ancestry shared by Omotic-speaking peoples is found predominantly in present-day southern Ethiopia and is associated with haplogroup E, thus revealing a plausible source.
Note that this is also consistent with the origin of anatomically modern humans in East Africa, as Ethiopia is located in East Africa.
The Omotic ancestry (indigenous African ancestry) of the Natufians being the source of their paternal haplogroup (E1b1b1b2-Z830) implies that E1b1b originated among indigenous Africans. Considering this and considering the predominance of indigenous Africans in regard to the overwhelming majority of Haplogroup E’s other subclades, it is logical to conclude that E1b1b originated among pure indigenous Africans.
Spread of E1b1 throughout africa
Given that it has been concluded herein that Paternal Haplogroup E remained in Africa when carriers of Paternal Haplogroups C, F, and D migrated out of Africa, there arises a curiosity regarding E’s travels throughout the continent during and after that time - particularly E1b1’s travels, since it is by the far the most populous subclade of E. The travel routes within Africa of males who possessed subclades of E since that time can be determined by considering the following:
East Africa as the original homeland of anatomically modern humans and therefore the original homeland of carriers of Paternal Haplogroups C, F, D and E.
The hypothesized migration routes that could have led carriers of Paternal Haplogroups C, F, and D out of Africa, with the presumption that carriers of E inhabited East Africa with them or traveled among them for some time until they ultimately diverged onto their own path.
Hypothesized Migration Routes of Population CFD
The population that eventually split into carriers of Paternal Haplogroups C, F, and D and carriers of Paternal Haplogroup E will be referred to by the aforementioned designation, Population CFDE. And the subsequent populations into which it split will be referred to by the aforementioned designations Population CFD and Population E, respectively.
There are two hypotheses regarding the route out of Africa via which Population CFD traveled: the Northern Dispersal Route Scenario and the Southern Dispersal Route Scenario. The former posits that Population CFD migrated out of Africa via Egypt, and the latter posits that they did so via a land bridge between the Horn of Africa (Eritrea and Djibouti) and Yemen.
Southern Dispersal Route Scenario: If Population CFD migrated out of Africa via a land bridge between the Horn of Africa and Yemen, then Population CFDE would have split into Population CFD and Population E while still in East Africa, likely Ethiopia (since that is the home of the oldest anatomically modern human fossils). Therefore, Population E would have traveled from Ethiopia to Sudan, then Chad, then Niger, then Mali, and possibly Mauritania; this path will be referred to as Route-A and is labeled as Population E’s Route (A) in the map below.
Northern Dispersal Scenario: If Population CFD migrated out of Africa via Egypt, then Population CFDE could have still split while in Ethiopia but it could have also split while as far north as Egypt.
So, if it split while in Ethiopia, then Population E would have likely traveled along the aforementioned Route-A.
However, if it split while as far north as Egypt, then Population E would have likely traveled through Libya, then possibly Tunisia, then Algeria, and finally Morocco; this path will be referred to as Route-B and is labeled as Population E’s Route (B) in the map below.
Alternatively, Population CFDE could have experienced two splits by males possessing Haplogroup E: one that was carried out by E-carrying males who traveled along Route-A and a second one that was carried out by E-carrying males who traveled along Route-B.
The third possibility is what likely occurred, due to the predominance of Paternal Haplogroup E1b1a (E-V38) along Route (A) and below - and the predominance of Paternal Haplogroup E1b1b along Route (B). The distribution of these two subclades of E and the deduced time of the migration out of Africa (50,000 YBP) indicate that there were two splits from Population CFDE and they were each comprised largely of males possessing E1b1 (E-P2), which is the direct ancestral subclade of E1b1a and E1b1b; E1b1 formed 53,000 YBP [Link] and therefore would have been carried by a considerable number of males by 50,000 YBP, but E1b1a and E1b1b did not form for another 8,600 years (41,400 YBP).
A map highlighting the paths that Population CFD and Population E likely traveled from East Africa to their destinations.
At 41,400 YBP, a male who descended from the E1b1-carrying males who split first and traveled along Route (A) would have produced a son with the mutation for E1b1a, and his male descendants would have gone on to become the most numerous males in the region, ultimately replacing the ancestral E1b1 lineage therein.
Likewise, also at 41,400 YBP, a male who descended from the E1b1-carrying males who split second and traveled along Route (B) would have produced a son with the mutation for E1b1b, and his descendants would have gone on to become the most numerous in the region, ultimately replacing the ancestral E1b1 lineage therein.
In regard to E1b1a forming and becoming dominant along Route (A), which includes Niger (part of which is in the Sahara), there is archeological evidence that supports this scenario: dental affinities between fossils of individuals who inhabited Niger’s portion of the Sahara 8000 years ago and modern West Africans (E1b1a carriers) indicate that West Africans' ancestors inhabited the Sahara millennia ago. These affinities are explained in the scientific article Tracing the “Bantu Expansion” From Its Source: Dental Nonmetric Affinities Among West African And Neighboring Populations [Link].
The following is a quote from the article. Note that "Bantu" refers to Niger-Congo peoples (which includes West Africans) and various so-called "Bantu" peoples who split from the former a few millennia ago, thus spreading to other parts of Africa via the Bantu Expansion [Link]. These peoples possess various subclades of E (via its subclade E-M2), one of which is E1b1a - which is by far the most common one among them.
"Bantu-speaking populations living throughout sub-Saharan Africa are relative new-comers to this vast area. Archaeological, linguistic, and other evidence (e.g., oral histories) revealed that their origins lie to the northwest…. One small sample from Niger (ca. 7600 BC) was used to explore the purported proto-Bantu Saharan origins. The remaining samples, dating to the Iron Age and historic periods, are from west, west-central (i.e., western Bantu), and central (eastern Bantu) Africa. After editing, data from 24 traits were compared using the mean measure of divergence to yield inter-sample phenetic distances. The following results were obtained: 1) the Niger sample’s affinity to west Africans supports the possibility that it is representative of proto-Bantu ancestors, 2) temporal affinities are apparent, in that older samples are more similar to one another, and 3) spatial divergence is evident in the form of an apparent cline from more to less dental complexity between west Africa and the other regions. All told, these results seem supportive of the hypothesized migrations."
An 8000-year-old skull from the portion of Niger that is located in the Sahara desert (left) and the skull of a modern person from the Congo (right). The photo is from Tracing the “Bantu Expansion” From Its Source: Dental Nonmetric Affinities Among West African And Neighboring Populations.
In regard to E1b1b forming and becoming dominant along Route (B) tens of millennia ago, the paternal haplogroup of ancient Iberomaurusian specimens (as already discussed herein) supports this scenario: the remains of several Iberomaurusian individuals who lived between 15,100 and 13,900 YBP at Taforalt - Morocco, which is located along Route (B), were analyzed and determined to possess Haplogroup E1b1b1a1 (a subclade of E1b1b). Additionally, in support of Haplogroup E originating in Africa (and therefore among indigenous Africans) rather than being brought to Africa by Eurasians, the DNA of the Iberomaurusians of Taforalt was determined to be 36.5% “Sub-Saharan African”; this portion of their DNA was determined to be very, very similar to that of modern West Africans (Niger-Congo and Bantu peoples), as already explained herein.
