Map of Nigeria, showing the location of the Iwo Eleru rock shelter and the Iwo Eleru skulls. (Harvati et al., 2011)
Sub-Saharan Africans have an unusual complex of dental features:
[…] compared to other world populations, Africans south of the Sahara Desert are distinct dentally — especially in their expression of nine high- and two low-frequency morphological features. This suite of traits was termed the “Sub-Saharan African Dental Complex” (SSADC); it includes the world’s highest occurrences of Bushman canine, two-rooted UP1, UM1 Carabelli’s trait, three-rooted UM2, LM2 Y-groove, LM1 cusp 7, LP1 Tom’s root, two-rooted LM2, and UM3 presence, and among the lowest occurrences of UI1 double shoveling and UM1 enamel extension. (Irish, 2011)
The two low-frequency traits appear to be “derived.” They seem to have developed in sub-Saharan Africa after modern humans began to spread to other continents. The other traits, however, are ancestral:
[…] the same nine high-frequency traits are also ubiquitous in the dentitions of extinct hominids and many extinct and extant non-human primates
[…] The presence and, indeed, prevalence (see next section), of high-frequency Sub-Saharan dental traits in fossil and recent hominoids—some of which are probably direct ancestors of modern humans, suggests they have been around for a long time. (Irish, 1998, pp. 87-88)
In addition to these traits, Irish (1998) mentions a low-frequency trait that seems likewise ancestral and specific to sub-Saharan Africans:
A final ancestral feature found with some regularity in Sub-Saharan Africans, relative to other modern groups, is polydontia. Numerous cases of extra incisors, third premolars, and fourth molars have been noted […] In one study (Watters, 1962) the incidence reached 2.5-3% in several hundred west Africans; many of the extra teeth were fully formed and erupted. “Typical” mammals exhibit three incisors and four premolars (Jordan et al., 1992). Polydontia is also found in living non-human primates […] (Irish, 1998, p. 88)
Why are these ancestral traits much more common in sub-Saharan Africans than in other humans? There are several possible reasons. One is that non-Africans began as a small founder group and thus lost much of the dental variability that still characterizes Africans. Another reason might be that natural selection favored new forms of dentition outside Africa, perhaps as a response to new food sources or new ways of preparing food.
But there’s a third possible reason: archaic admixture. Just as modern humans mixed to some extent with Neanderthals in Europe and Denisovans in Asia, perhaps there was also mixture with archaic hominins in Africa, and perhaps this admixture introduced archaic dental features into present-day Africans.
But how could present-day Africans have archaic admixture? If modern humans originated in Africa, wouldn’t they have encountered archaic humans only in Europe and Asia?
Well, at first, modern humans did not occupy all of Africa. They were initially a small population somewhere in East Africa. Then, around 80,000 years ago, this population began to expand northward and eventually into Eurasia (Watson et al., 1997). Meanwhile, the same expansion was taking modern humans westward and southward into other parts of Africa.
Just whom exactly did these modern humans encounter during their expansion within Africa? Initially, they probably met hominins who looked the same but still lacked some of the mental rewiring that gave modern humans a competitive edge. These “almost-moderns” account for about 13% of the current sub-Saharan gene pool and may have been related to the Skhul-Qafzeh hominins who occupied the Middle East 120,000 to 80,000 years ago (Watson et al., 1997).
As modern humans spread further west and south within Africa, they encountered much more archaic hominins, and perhaps even lingering Homo erectus groups. About 2% of the modern African genome comes from an archaic population that split from ancestral modern humans some 700,000 years ago. This admixture is dated to about 35,000 years ago and may have occurred in Central Africa, since the level of admixture is highest in pygmy groups from that region (Hammer et al., 2011).
A more tangible sign of admixture is visible in a skull retrieved from the Iwo Eleru rock shelter, in southwestern Nigeria, and dated to approximately 16,300 BP:
Our analysis indicates that Iwo Eleru possesses neurocranial morphology intermediate in shape between archaic hominins (Neanderthals and Homo erectus) and modern humans. This morphology is outside the range of modern human variability in the PCA and CVA analyses, and is most similar to that shown by LPA individuals from Africa and the early anatomically modern specimens from Skhul and Qafzeh.
[… ] the transition to anatomical modernity in Africa was more complicated than previously thought, with late survival of “archaic” features and possibly deep population substructure in Africa during this time. (Harvati et al., 2011)
Then there is the Broken Hill skull, found near Kabwe, Zambia and dated to 110,000 BP (Bada et al., 1974). It looks for all the world like a Homo erectus. Textbooks generally try to raise it to Homo sapiens status or argue for an earlier dating. Recently, a late dating has been confirmed by Stringer (2011).
Interestingly, when Irish (2011) compared dentitions from west, central, east, and south Africa, ranging in age from the late Pleistocene to the mid-1950s, the early Holocene Kenyans and Tanzanians were the sample that had the fewest ancestral traits of the Sub-Saharan African Dental Complex (SSADC). In other words, the SSADC seems to have been least present in the “homeland” of modern humans (East Africa) and more present farther west and south.
Given the high level of archaic admixture in sub-Saharan Africans, we may have to revise downwards the estimate of 1 to 4% Neanderthal admixture in Eurasians. Yes, Eurasians are closer than sub-Saharan Africans to the Neanderthal genome. But is this discrepancy solely due to Neanderthal admixture in Eurasians? Could it also be due to Sub-Saharan Africans becoming further removed from the Neanderthal genome through admixture with other archaic groups?
The past may be a stranger country than previously thought. When farming villages began to form in the Middle East, there may still have been archaic hominins roaming over parts of western and southern Africa.
Bada, J.L., R.A. Schroeder, R. Protsch, & R. Berger. (1974). Concordance of Collagen-Based Radiocarbon and Aspartic-Acid Racemization Ages, Proceedings of the National Academy of Sciences (USA), 71, 914-917.
Hammer, M.F., A.E. Woerner, F.L. Mendez, J.C. Watkins, and J.D. Wall. (2011). Genetic evidence for archaic admixture in Africa, Proceedings of the National Academy of Science (USA), 108, 15123-15128, www.pnas.org/cgi/doi/10.1073/pnas.1109300108
Irish, J.D. (2011). Afridonty: the “Sub-Saharan African Dental Complex” revisited, American Journal of Physical Anthropology, 144(supp. 52), 174
Irish, J.D. (1998). Ancestral dental traits in recent Sub-Saharan Africans and the origins of modern humans, Journal of Human Evolution, 34, 81-98.
Harvati, K., C. Stringer, R. Grün, M. Aubert, P. Allsworth-Jones, C.A. Folorunso. (2011). The Later Stone Age Calvaria from Iwo Eleru, Nigeria: Morphology and Chronology. PLoS ONE 6(9): e24024. doi:10.1371/journal.pone.0024024
Stringer, C. (2011). The chronological and evolutionary position of the Broken Hill cranium. American Journal of Physical Anthropology, 144(supp. 52), 287
Watson, E., P. Forster, M. Richards, and H-J. Bandelt. (1997). Mitochondrial footprints of human expansions in Africa, American Journal of Human Genetics, 61, 691-704.