Tuesday, May 11, 2010

2010 Berber mtDNA study finds Berber roots foundational in Africa - Frigi 2010

Ancient Local Evolution of African mtDNA Haplogroups in Tunisian Berber Populations

Frigi et al.

Human Biology (August 2010 (82:4)

Abstract

Our objective is to highlight the age of sub-Saharan gene flows in North Africa and particularly in Tunisia. Therefore we analyzed in a broad phylogeographic context sub-Saharan mtDNA haplogroups of Tunisian Berber populations considered representative of ancient settlement. More than 2,000 sequences were collected from the literature, and networks were constructed. The results show that the most ancient haplogroup is L3*, which would have been introduced to North Africa from eastern sub-Saharan populations around 20,000 years ago. Our results also point to a less ancient western sub-Saharan gene flow to Tunisia, including haplogroups L2a and L3b. This conclusion points to an ancient African gene flow to Tunisia before 20,000 years BP. These findings parallel the more recent findings of both archaeology and linguistics on the prehistory of Africa. The present work suggests that sub-Saharan contributions to North Africa have experienced several complex population processes after the occupation of the region by anatomically modern humans. Our results reveal that Berber speakers have a foundational biogeographic root in Africa and that deep African lineages have continued to evolve in supra-Saharan Africa.

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Ancient Local Evolution of African mtDNA Haplogroups in Tunisian Berber Populations
Frigi et al. Human Biology August 2010 (82:4)

Discussion
In this study we attempted to better elucidate the ancient African genetic background in the northwest African area, particularly in Tunisia. To this aim, we focused our study on Berber populations that are considered representative of the ancient North African populations that probably derived from Neolithic Capsians. During historic times, Berbers experienced a long and complicated history with many invasions, conquests, and migrations by Phoenicians, Romans, Vandals, Byzantines, Arabs, Bedouins, Spanish, Turks, Andalusians, sub-Saharans (communities settled in Jerba and Gabes in the 16th–19th centuries), and French (Brett and Fentress 1996). During these invasions, Berbers were forced back to the mountains and to certain villages in southern Tunisia (Fadhlaoui-Zid et al. 2004). At present, they are restricted to some isolates in the south who maintain the Berber language and to some populations in the north who lack an origin language. Many genetic studies on Tunisian Berber populations demonstrate the heterogeneity of Berbers with respect to European and sub-Saharan African contributions and the mosaic structure of Tunisian Berber populations with an absence of ethnic, linguistic, and geographic effects (Cherni et al. 2010).

In the present work, mtDNA data show a diversified distribution of African haplogroups. However, a question remains concerning the date of the sub-Saharan African inputs. Our results demonstrate an ancient local evolution in Tunisia of some African haplogroups (L2a, L3*, and L3b). The most ancient haplogroup is L3*, which would have been introduced from eastern sub-Saharan populations to North Africa about 20,000 years ago. The Siwa oasis sample studied by Coudray et al. (2009) contains sub-Saharan haplogroups L0a1, L3i, L4*, and L4b2, which are different from our Tunisian samples, in agreement with the heterogeneity of Berbers already shown in Tunisia.

Stevanovitch et al. (2004) suggested that the Gurna population in Egypt has conserved the trace of an ancestral genetic structure from an ancestral East African population characterized by a high haplogroup M1 frequency. This haplogroup is also present in three Berber populations (Kesra, Matmata, and Sned) with variable frequencies. In each of these populations, haplogroup L3* is also present. The association of both eastern African haplogroups in the Berber populations is a strong argument in favor of eastern African gene flow in Berbers. Other genetic and archaeological studies confirmed the crucial idea that an ancient population in East Africa constituted the basis of the ancestors of all African Upper Paleolithic populations—and their subsequent present-day descendants (Bengtson 2008; Keita 2004; Relethford 2000; Zakrzewski 2003, 2007).

Moreover, Berber languages spoken exclusively by North African populations belong to the Afro-Asiatic language. Diakonoff (1998) showed an exclusively African origin (Diakonoff, 1981, 1988) for the family. He explicitly described proto-Afro-Asiatic vocabulary as consistent with non-food-producing vocabulary and linked it to pre-Neolithic cultures in the Levant and in Africa south of Egypt. Moreover, Ehret. (2003) suggested that early Afro-Asiatic languages were spread by Mesolithic foragers from Africa into the Levant. On the contrary, Diamond and Bellwood (2003) suggested that food production and the Afro-Asiatic language family were brought simultaneously from the Near East to Africa by demic diffusion—in other words, by a migration of food-producing peoples. The evidence presented by Wetterstrom (1993) does not support this latter suggestion, however, and indicates that early African farmers in the Fayum initially incorporated Near Eastern domesticates into an existing indigenous foraging strategy and only over time developed a dependence on horticulture.

In conclusion, the crucial linguistic finding is that the three deepest clades of the Afro-Asiatic family are localized in Eritrea and Ethiopia. All the other languages of the family outside that region belong to subclades of just one of those deep clades. This kind of cladistic distribution is a basic criterion of the genetic argument for the genetic lineage origins well understood by geneticists. It applies to linguistic history as well.

Our results also point to a less ancient western African gene flow to Tunisia involving haplogroups L2a and L3b. Thus the sub-Saharan contribution to northern Africa starting from the east would have taken place before the Neolithic. The western African contribution to North Africa should have occurred before the Sahara’s formation (15,000 years BP). It seems likely that an expansion would have taken place in the Sahel zone starting about the time of a gradual climatic return to wetter conditions, when the Senegal River cut through the dunes (Burke et al. 1971). For subhaplogroup L2a1 (data not shown) we found some haplotypes that the Tunisian Berbers shared with Mauritanians and western sub-Saharan populations speaking a Niger-Congo language (studied by Salas et al. 2002).

This suggests that the people who brought these markers to the Berber populations most likely came from West African populations that spoke languages belonging to the Niger-Congo family when the Sahara became drier. However, this contribution of West African haplotypes and of other haplotypes, such as those belonging to haplogroup L1b1, could have been introduced to North Africa more recently.

Indeed, this West African contribution was difficult to date, because few haplotypes belonging to western African haplogroups have been observed, most of them being divergent. This result can be interpreted in different ways. Ancient western African mtDNA contributions could have disappeared from North Africa as a result of recent flows, or the situation observed now could be the result of a strong drift effect on ancient western African lineages, particularly those belonging to haplogroups L2a and L3b. A strong Iberian gene flow may have contributed to the decrease in African haplogroups. Indeed, most of the older hypotheses about North African population settlement used to suppose an Iberian or an eastern origin. The dates for subhaplogroups H1 and H3 (13,000 and 10,000 years, respectively) in Iberian and North African populations allow for this possibility. Kefi et al.’s (2005) data on ancient DNA could be viewed as being in agreement with such a presence in North Africa in ancient times (about 15,000–6,000 years ago) and with the fact that the North African populations are considered by most scholars as having their closest relations with European and Asian populations (Cherni et al. 2008; Ennafaa et al. 2009; Kefi et al. 2005; Rando et al. 1998).

However, considering the general understanding nowadays that human settlement of the rest of the world emerged from eastern northern Africa less than 50,000 years ago, a better explanation of these haplogroups might be that their frequencies reflect the original modern human population of these parts of Africa as much as or more than intrusions from outside the continent. The ways that gene frequencies may increase or decrease based on adaptive selection, gene flow, and/or social processes is under study and would benefit from the results of studies on autosomal and Y-chromosome markers.

Since the end of the extreme Saharan desiccation, lasting from before 25,000 years ago up to about 15,000 years ago, the Sahara has had post- and pre- Holocene cyclical climatic changes (Street and Grove 1976), and corresponding increases and decreases in population are probable. Wetter phases with better habitats perhaps allowed for increased colonization and gene and cultural exchange. Desiccation would have encouraged the emigration and segmentation of populations, with resultant genetic consequences secondary to drift producing more variation. During the last glacial period, the Sahara was even bigger than it is today, extending south beyond its current boundaries (Ehret 2002). About 13,000 years ago, large parts of the Sahara were as dry as the desert is now (White and Mattingly 2006). The end of the glacial period brought more rain to the Sahara, especially from about 8500 to 6000 BC (Fezzan Project 2006). By around 3400 BC, the monsoon retreated south to approximately where it is today, leading to the gradual desertification of the region (Kröpelin 2008). Thus the Sahara, through its cyclical environmental changes, might be seen as a microevolutionary “processor” and/or “pump” of African people that “ejected” groups to the circum-Saharan regions in times of increasing aridity.

Indeed, it must be noted that the high frequencies of cDe, P, and V antigens and low frequencies of FY antigens in some Berber-speaking groups (Chamla 1980; Mourant et al. 1976) indicate affinities with tropical Africans. These data may indicate recent or ancient gene flow from sub-Saharan Africa, a common immediate pre-Holocene ancestral group, or chance resemblance.

Our findings are in accordance with other studies on Y-chromosome markers that have shown that the predominant Y-chromosome lineage in Berber communities is the subhaplogroup E1b1b1b (E-M81), which emerged in Africa, is specific to North African populations, and is almost absent in Europe, except in Iberia (Spain and Portugal) and Sicily. Molecular studies on the Y chromosome in North Africa are interpreted as indicating that the southern part of Africa, namely, the Horn/East Africa, was a major source of population in the Nile Valley and northwest Africa after the Last Glacial Maximum, with some migration into the Near East and southern Europe (Bosch et al. 2001; Underhill et al. 2001). Hence, contrary to the suggestion that mtDNA haplogroups were introduced mostly from Iberia, it seems that Y-chromosome markers have an eastern African origin with an ancient local evolution in North Africa. These observations are in agreement with the proposal that the ancient communities ancestral in language to more recent Berber communities absorbed a lot of females from the existing pre-Holocene populations. This would indicate that the North African populations arose from admixture rather than from local evolution, leading to an intermediate genetic structure between eastern sub-Saharan Africans and Eurasians. Rock paintings in North Africa that show people of different phenotypes living together are a strong argument for our hypothesis (Hachid 1982, 1992, 1998).

In conclusion, our findings parallel the more recent findings of both archaeology and linguistics on the prehistory of Africa. The present study suggests that sub-Saharan contributions to North Africa have experienced several complex population processes after the occupation of the region by anatomically modern humans. Our results reveal that Berber speakers have a foundational biogeographic root in Africa and that deep African lineages have continued to evolve in supra-Saharan Africa.

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Biocultural Emergence of the Amazigh in Africa: Comment on Frigi et al. (2010)
S. O. Y. Keita. Human Biology (August 2010) (82:4)

Frigi et al. [2010 (this issue)] present some new findings on a population of Amazigh—Berber speakers in Tunisia. Although their study is not exhaustive, they provide an outline of human population history in the Maghreb and a general discussion of its mtDNA diversity. Their work is important in inviting researchers to think about the concepts of continuity and change in biology, culture, language, and identity in a geographic space. Their presentation helps in understanding the complexity of examining the ancestry and emergence of Berber origins in Africa as a local process and encourages the consideration of many questions about how the human biology and culture of a known population or ethnolinguistic group can be conceptualized through space and time.

Berber- (Tamazight-) speaking communities are thought to represent the clearest known descendants of the ancient indigenous populations of Africa west of the northern Nile valley in the supra-Saharan and northern Saharan regions (Brett and Fentress 1996; Camps 1982; Desanges 1981). “Indigenous” here can refer only to those whom we can perceive as having had the longest tenure on the land, using available historical evidence. However, there are questions. What constitutes “historical evidence” for earlier periods? Should it include archaeology, paleontology, historical linguistics, skeletal biology, and genetics, as broadly advocated by a historical anthropological approach (e.g., Kirch and Green 2001; Mace et al. 2005)? Or is it only to be based on the interpretation of texts from the ancient Egyptian, Greco-Roman, or Islamic periods [e.g., see comments by Brett and Fentress (1996), Desanges (1981), Norris 1982, and Snowden (1971)]?

How is the varied evidence to be ranked in importance, reconciled when it seems to be in contradiction, and analyzed synthetically? A simplistic positivism has to be avoided in discussions of any facet of human history because various pathways could lead to similar results. What is the role of evolutionary mechanisms—adaptive selection, gene flow, drift, sexual selection—in explaining the biology of some of the ancestors of Berber speakers at the deepest time levels and of living Amazigh as well? Frigi et al. (2010) do not address all these questions directly, but their work implicitly acknowledges their importance and provides a new framework for investigation.

Braudel’s (1980) concept of different levels of history can be adapted and adopted in a modified form as levels of biocultural or bioethnic history, to further consider Frigi and colleagues’ contribution, which implicitly acknowledges the contingent and multidimensional character of population interactions through time against an evolutionary background.

Another issue of some interest is the (mis)labeling of Berbers as “Eurasian” migrants from the Near East: Did they arrive as a unit from Asia or Europe, as a settler colonist “package” with a persistent identity analogous to Europeans in South Africa, or did the biology, language, and culture of the Amazigh emerge primarily from a set of interactions in Africa involving African peoples at base, that is, as a part of authentic African historical and biological processes? There are no ancient Berber communities outside Africa, and the idea of simple demic diffusion of Berbers as a people to the Maghreb (e.g., Arredi et al. 2004) from the Near East is not supported. It is of some interest that even Coon and Hunt (1965), using a raciotypological paradigm now long discredited, postulated a massive invasion of Africa by “Caucasians” in the Pleistocene and therefore thought that Berber language and identity had entered the Maghreb from more southerly regions in Africa. Frigi and colleagues suggest that several populations over time were involved in the biological ancestry of the current Berber speakers, and this is consistent with archaeological evidence of actual migration in the mid- to late Holocene (Camps 1982) as well as historical documentation. Craniofacial diversity has been documented in the region before Vandal and Arab migrations (Keita 1990).

It is important to remember that biology, language, and culture are not intrinsically or obligatorily correlated, a principle established some time ago (Boas 1940). It is not particularly surprising that one can sometimes find markers that will correlate across biology and culture at some levels—but the issue is how this came to be and when, and what it represents historically and socially. There is always the question of whether the correlation is an artifact of recent events or of “primordial” ontology. A language or language family can be adopted slowly or rapidly by nonnative speakers. The current biological profile of a region may predate (or postdate) the language spoken there. Communities may adopt (nonlinguistic) cultural practices from others without greatly changing language or biology; or they may become primarily integrated linguistically and politically but not biologically or exhibit other permutations of these variables, such as total biocultural assimilation. The biology of a particular ethnolinguistic group or community may change based on intermarriage if the social rules allow the offspring to become group members. Such matings may have occurred long before the recall of communal memory, whether in texts or oral tradition. Relatively nonethnocentric polygamous societies or populations may have cultural descendants who are genealogically heterogeneous when viewed over a millennium.

Frigi et al. (2010) suggest these possibilities as factors in their consideration of the asymmetric assimilation of females of non-African origin into Berber-speaking populations whose males currently have a predominance of lineages defined by the African M35/81 biallelic marker. It is interesting that these “non-African” mtDNA lineages are usually predominant while being diverse (Coudray et al. 2009; Fadhlaoui-Zid et al. 2004; Khodjet-el-Khil et al. 2008). The existence of mtDNA lineages common to Saami and some Amazigh groups (Achilli et al. 2005) is likely to be explained by the migration of females bearing these lineages from a region in northern Europe (perhaps in the ranks of the Vandals or far more ancient back-migrations to Africa), whether they were ethnically Saami or not. However, there may have been other locales where these lineages once existed. Circular reasoning in syntheses involving multiple disciplines has to be avoided. The criteria and methods for a given discipline usually have to be given equal weight, and their results should be considered independently before an effort at synthesis is made. For example, a hypothesis about the place of origin of a language family or phylum must be based on linguistic evidence and methods, not on DNA or craniofacial patterns. Likewise the place of origin of a particular genetic variant or lineage has to be based on genetic data, principles, and models, not on archaeological data. The locale of origin of a particular culture or archaeological industry is subject to analyses based on methods and theory that are specific to the relevant disciplines.

The only exception to these “rules” is if a calculated date of origin of a genetic variant found in a given locale predates the existence of people in that place. Although the notion of population ties together both biology and culture broadly conceived, it cannot be claimed that continuity in one necessarily means continuity in another. If the question is about physical population migration, then the same conclusion reached from every discipline independently would seem to best support the claim (Rouse 1986). However, it cannot be said absolutely that there was no movement if all lines of evidence do not point in the same direction. The idea of Occam’s razor may sometimes mean accepting the reality of human complexity and an inability to reconcile evidence with preconceived models.

Frigi et al. (2010) have provided a general temporal framework for Maghreb population history, from the Paleolithic to French colonization. This is appropriate given the evidence for early modern human behavior and life history in the Maghreb (Bouzouggar et al. 2007; T. Smith et al. 2007), the diversity of various epi-Paleolithic and Neolithic cultures in or near the region associated with climatic changes (Lubell et al. 1984; Rahmani 2003, 2004; Sereno et al. 2008; Sheppard and Lubell 1990), and the interactions with known “peoples” at later dates (Bennett 1960; Brown 1968; Desanges 1981; Hirschberg 1960; Nebel 2002).

In reviewing data from multiple disciplines, Frigi and colleagues have given the region’s populations a multidimensional existence. In providing evidence for the ongoing microevolution in the Maghreb of ancient mtDNA lineages that emerged in Africa and evidence of later gene flow from multiple directions, they have revealed that this region has biological continuity with the deep past as well as change.

Frigi and colleagues may have inadvertently revealed peoples whose ancestors had a level of cultural flexibility in accepting outsiders as mates. As noted, the male and female histories of a population may be different in their sources (Wilkins 2006), although they are now seen as part of a recognizable biocultural entity with a categorically singular identity. How is the emergence of the Amazigh peoples in the geographic range of their homeland to be understood in terms of culture, language, and biology? In some sense the question is about origins, a term that can be confusing because of its various meanings. It can be applied to different aspects of a population—which can be disarticulated and can change as a function of time. Ancestry must not be confused with explanation, or gene history with population or culture history. Known ancestors and the “ancestors of one’s genes” are not the same things necessarily (Weiss and Long 2009).

Can a narrative of “origins” be constructed on the basis of an internal perspective of the dynamics of the human communities of northwestern Africa, considered through time? Or is this region simply an appendage of other places? Sometimes the Amazigh, by use of the term “Eurasian” in a categorical model of analysis, are placed in a raciotypological model without reference to evolution and their indigenous emergence in Africa. (It can be ventured that this is largely based on nonevolutionary ideas about phenotype, notions of bounded unchanging populations, problematic assumptions about language families, and certain old attitudes and theories about Africa.)

Frigi and colleagues have documented the deep-time biological connections of current Berber speakers to Africa. The migration of “Europeans” and “Asians” is also discussed. There has been continuity and change in the population from original settlement. It is important to remember that high levels of gene flow and biocultural assimilation could lead to great biological heterogeneity in a population whose language family or culture does not change. Frigi and colleagues address the idea of the indigenous, although not explicitly, and lay the groundwork for more nuanced future discussions. They suggest a complex biogeographic history not reducible to raciotypological constructs or outdated simplistic theories of colonization and migration. They provide a basis for a rich discussion by acknowledging the interactions of known peoples in the Maghreb and unknown actors of a deeper past.

The issue of what is indigenous is seen to be one of definition, turning on what aspect of a population or region is ranked as its “defining” characteristic, and whether this may change or could have changed over time. The term indigenous unfortunately is connected to a discourse about the West and non-West and sometimes has a negative sensibility and hence may not be the best word, but a discussion of this issue is beyond the scope of this presentation. Of course “indigenous” is a relative term when the temporal scope of human evolution and history is considered, and it even seems to depend to a degree on what part of the world is under discussion. Europe can serve as a good example. If it is asked who are the “indigenous” Europeans, there would probably be a request to clarify the time depth, given that modern humans are not native to Europe and arrived there from elsewhere.

(The next question therefore is at what point do they become “European” and what precisely does this mean: current limb proportions, skin color, genetic variation, language, the presence of Neanderthal DNA?) Does “indigenousness” require residency back to the upper Paleolithic, the Neolithic, and so on? Is it only a biological phenomenon requiring a “drop” of Neanderthal blood or a linguistic phenomenon requiring the speaking of Indo-European languages? Or if the question is who were the indigenous inhabitants of northern, southern, western, eastern, or central Europe, the answers would necessarily take on a different tone, based on other information.

Are the Basque speakers the indigenous inhabitants of Europe, if currently spoken language phyla and families are used as “population markers,” a problematic assumption? Basque predates Indo-European, and there is some indication of some level of biological distinctiveness (Alonso et al. 2005). The fact that historical linguists (e.g., Ehret 2002; Nichols 1997) can reconstruct the existence of culture-linguistic units for a proto-language family (e.g., Adamawa) or phylum (e.g., Niger-Congo), which may have migrated, does not mean that they are suggesting that the people making up such entities connote genetic units or Mendelian breeding populations. It also does not mean that the speakers of such proto-entities had a common molecular or social genealogical origin at foundation, or that the linguists are suggesting this.

Defining “origins” or “indigenous” becomes one of perspective. How much “Basque ancestry” would a European population have to have for the label of “indigenous European” to apply? If none, why not? (What is the relationship between cultural and biological genealogy?) Can it be assumed that the Basques of today biologically represent those of the past “accurately”? The post-Paleolithic European assimilation of males from Africa and Asia bearing younger genetic variants is documented (Cruciani et al. 2004, 2007): Are such ancient admixed populations to be viewed as “less” European or non-European? Are Nordics or the Basques the “standard” European? Is language, biology, culture, geography, or something else the arbiter of European-ness? In practice, this question seems to be little asked in studies of Europeans: All these groups and nationalities are considered European with little question. Aegean peoples are not presented as “hybrids.” The linguistic and genetic diversity is not a factor in the designation of “indigenous” for Europe.

But in the case of Africa there seems to be a problem with diversity for some scholars. The Indo-European language phylum, in the standard evidence-based interpretation, did not originate in the European heartland (Ehret, personal communication, 2010). Most people in Europe today speak Indo-European languages—now considered as “indigenous” as Basque. What does it mean for the concept of European if Europe’s major language phylum did not originate in what is considered Europe proper? How much of the spread of early Indo-European was due to outright settler colonization and how much to language shift—these are questions that will likely be debated for some time. Are the Finns, Saami, and Hungarians (or their “original” ancestors)—all non-Indo- European-speaking—to be considered Europeans? Apparently so. Contrast this with ideas held by some about Berbers as “Eurasians” who speak a language family that belongs to a phylum whose proto-parent emerged in Africa using standard historical linguistic criteria and whose major history and differentiation occurred in Africa (Ehret 2002; Greenberg 1963; Nichols 1997). In discussions about Europe, geography seems to be enough to define what is “indigenous”— with the exception of the Turks. This contrast deserves review by students of the sociology of knowledge.

The European example is relevant to the discussion of Berbers because of the use of terms by some researchers that imply that Berbers are not an African development, an African people, in their beginning and current state. Calling the Amazigh “Eurasian” based primarily on skin color without a discussion of process in history, language, evolution, and Y-chromosome variants can easily be seen as problematic when literature about Europe is examined carefully. The possibility of asymmetric gene flow with more Eurasian females being assimilated into the ancient Maghreb—and their lineages simply differentially surviving in greater frequencies—is addressed in a preliminary fashion by Frigi et al. (2010) and further engages us in the history of social interactions that may influence population biology.

Returning to the Amazigh, the findings and comments of Frigi et al. (2010) on Tunisian Berbers, and Berbers in general, suggest a new way of looking at the Maghreb region of Africa. Their review and analysis offer the opportunity to begin to develop a new and nuanced narrative about the peopling of the region, one that avoids the biases of past writings. Their findings of ongoing evolution in the Maghreb of ancient mtDNA lineages that originated in Africa, synthesized with the evidence of the assimilation of migrants (mainly female?) from Europe and the Near East, the predominance of uniquely African Y-chromosome lineages, and the observation that Berber is the only extant indigenous language in the region suggest the workings of both biological and cultural processes.

There are clearly different levels of biological and cultural history. Except in situations of migrationist settler colonialism associated with the annihilation or conquest of local peoples, groups emerge from local elements and new additions—all influenced by the social and physical environments. This view of populations as assemblages and processes is different from a notion of them as essentialist primordial entities with fixed traits having continuity over time. In any geographic space groups can interact at various levels with various strictures; languages can be adopted partly or fully, and social rules may allow the acceptance of offspring by foreign females but not males, or vice versa. It is possible for a group to view itself as genealogically homogeneous by memory but to evince a genetic heterogeneity of lineages obtained in the remote past. “Admixture” in the late Pleistocene in the deep background of a regional population is to be differentiated from gene flow between known historical entities.

Frigi and colleagues’ suggestion that supra-Saharan Africans are an indigenous development with a complex story is a corrective to past models. The settlement of the coastal Maghreb in the Middle and Late Stone Age is a part of the settlement of the world outside Saharo-tropical Africa. The early modern human presence in the Maghreb suggests that that region played a role in modern human developments (Bouzouggar et al. 2007). Even if whole communities later came from outside Africa into the Maghreb (before the Phoenicians), which is not knowable, they became thoroughly assimilated into the autochthonous population— which adopted some of its culture (Camps 1982), and their descendants are a part of the emergence of the much later Amazigh world.

Less arid climatic conditions in the early to mid-Holocene Sahara allowed for the interaction of various peoples who no doubt contributed to the population history, as observed by Frigi et al. (2010). Saharan developments likely help to explain the Berber emergence, because, based on recent work [see Kuper and Kropelin (2006) and Sereno et al. (2008)], the desert was likely the site of a metapopulation and cultural differentiation. Whether the early Saharan rock paintings depict only Africans of varying phenotypes or such Africans and Asians (as suggested by Frigi and colleagues) can be debated, but the net result was assimilation into Amazigh communities, because there are no Berbers in Europe or Asia. The light skin color of Mediterranean Africa may be the result of adaptive evolution or drift, given the length of time of modern people in Africa, including the Maghreb, or gene flow, but more likely some combination.


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Elite white universities discriminate against Asians using reverse "affirmative action"
http://nilevalleypeoples.blogspot.com/2014/06/elite-white-universities-discriminate.html

Deteriorating state of white America
http://nilevalleypeoples.blogspot.com/2014/05/deteriorating-state-of-white-america.html


Racial Cartels (The Affirmative Action Propaganda machine- part 2
http://nilevalleypeoples.blogspot.com/2014/05/the-affirmative-action-propaganda.html

Hereditarian's/HBD's "Great Black Hope"
http://nilevalleypeoples.blogspot.com/2014/04/blog-post.html



Exploding nonsense: the 10,000 Year Explosion
http://nilevalleypeoples.blogspot.com/2014/03/exploding-nonsense-review-of-cochran_8.html


We need "rational racism"- Convicted felon Dinesh Dsouza becomes his own test case
http://nilevalleypeoples.blogspot.com/2014/01/we-need-rational-racism-proponent.html

The Affirmative Action Propaganda Machine- part 1
http://nilevalleypeoples.blogspot.com/2014/01/the-affirmatve-action-propaganda.html

Two rules for being "really" black- no white wimmen, no Republican
http://nilevalleypeoples.blogspot.com/2014/01/to-be-really-black-you-cant-have-white.html

The Axial age reconsidered
http://nilevalleypeoples.blogspot.com/2014/01/the-axial-age-reconsidered.html

Cannibal seasonings: dark meat on white
http://nilevalleypeoples.blogspot.com/2013/12/i.html

"Affirmative Action" in the form of court remedies has been around a long time- since the 1930s- benefiting white union workers against discrimination by employers
http://nilevalleypeoples.blogspot.com/2013/09/affirmative-action-as-term-appears-in.html

Mugged by reality 1: White quotas, special preferences and government jobs
http://nilevalleypeoples.blogspot.com/2013/06/mugged-by-reality-1-white-quotas.html


Lightweight enforcement of EEO laws contradicts claims of "flood" of minorities "taking jobs"
http://nilevalleypeoples.blogspot.com/2013/06/blog-post.html

Railroaded 3: white violence and intimidation imposed quotas
http://nilevalleypeoples.blogspot.com/2013/06/railroaded-3-white-violence-and.html

Railroaded 2: how white quotas and special preferences blockade black progress...
http://nilevalleypeoples.blogspot.com/2013/06/railroaded-2-thow-white-quotas-and.html

Railroaded 1: How white affirmative action and white special preferences destroyed black railroad employment...
http://nilevalleypeoples.blogspot.com/2013/06/railroaded-how-white-affirmative-action.html

Affirmative action: primary beneficiaries are white women...
http://nilevalleypeoples.blogspot.com/2011/04/quick-regime-kill-hopes-in-libya.html

7 reasons certain libertarians and right-wingers are wrong about the Civil Right Act
http://nilevalleypeoples.blogspot.com/2012/05/7-reasons-libertarians-may-be-wrong.html

Assorted "Role models" debunked
http://nilevalleypeoples.blogspot.com/2009/11/blog-post_5570.html


Social philosophy of Thomas Sowell
http://nilevalleypeoples.blogspot.com/2011/07/social-philosophy-of-thomas-sowell.html


Additional gene flow data... :)


Bogus "biodiversity" theories of Kanazawa, Ruston, Lynn debunked
http://nilevalleypeoples.blogspot.com/2010/09/blog-post.html

JP Rushton, Michael Levin, Richard Lynn debunked. Weaknesses of Jared Diamond's approach. 
http://nilevalleypeoples.blogspot.com/2010/04/blog-post_1818.html

In the Blood- debunking "HBD" and Neo-Nazi appropriation of ancient Egypt
http://nilevalleypeoples.blogspot.com/2009/11/blog-post_29.html

early Europeans and middle Easterners looked like Africans. Peoples returning or "backflowing" to Africa would already be looking like Africans
http://nilevalleypeoples.blogspot.com/2010/05/blog-post_1754.html

 Ancient Egypt: one of the world's most advanced civilizations- created by tropical peoples
http://nilevalleypeoples.blogspot.com/2010/09/blog-post_06.html

Playing the "Greek defence" -debunking claims of Greeks as paragons of virtue or exemplars of goodness
http://nilevalleypeoples.blogspot.com/2013/03/playing-greek-defence-review-of-thornton.html

Quotations from mainstream academic research on the Nile Valley peoples
http://nilevalleypeoples.blogspot.com/2010/04/blog-post_9251.html


http://egyptsearchreloaded.proboards.com/thread/15/basic-database-nile-valley-studies



OTHER LINKS
Race, IQ, and Wealth: What the facts tell us about a taboo subject By Ron Unz


HBD EVOLUTION, BRAIN SIZE AND NATIONAL IQ CLAIMS DEBUNKED
Evolution, brain size, and the national IQ of peoples ... - Jelte Wicherts 2010
http://wicherts.socsci.uva.nl/wichertsPAIDrejoinder.pdf
------------------------------------

Why national IQs do not support evolutionary theories of intelligence - WIcherts, Borsboom and Dolan 2010
Personality and Individual Differences 48 (2010) 91-96
http://wicherts.socsci.uva.nl/wicherts2010.pdf
----------------------------- -------------

Are intelligence tests measurement invariant over time? by JM Wicherts - ?2004
 --Dolan, Wicherts et al 2004. Investigating the nature of the Flynn effect. Intelligence 32 (2004) 509-537
http://www.iapsych.com/iqmr/fe/LinkedDocuments/wicherts2004.pdf
-------------------------------------------

LYNN AND VANHAVEN'S IQ AND THE WEALTH OF NATIONS DEBUNKED
---------------- -------

www.anth.uconn.edu/faculty/mcbrearty/Pdf/McB%20&%20Brooks%202000%20TRTW.pdf

------------------------

Race and other misadventures: essays in honor of Ashley Montagu... By Larry T. Reynolds, Leonard Lieberman

http://books.google.com/books?id=5DLrgG_MflgC&pg=PA190&dq=r-+k-+selection+races&cd=1#v=onepage&q=r-%20k-%20selection%20races&f=false
--------------------------------

Race and intelligence: separating science from myth. By Jefferson M. Fish. Routledge 2002. See Templeton's detailed article referenced above also inside the book

http://books.google.com/books?id=t9OdPPLIgMAC&pg=PA64&dq=r-+k-+selection+races&cd=7#v=onepage&q=r-%20k-%20selection%20races&f=false
------------------------

http://www.ogiek.org/indepth/what-they-mean.htm
---------------- -------

Oubre, A (2011) Race Genes and Ability: Rethinking Ethnic Differences, vol 1 and 2. BTI Press
For summary see: http://www.skeptic.com/eskeptic/05-02-18/
---------------- -------

http://www.dartmouth.edu/~chance/course/topics/curveball.html

-----------------------------------------------------------

--S OY Keita, R A Kittles, et al. "Conceptualizing human variation," Nature Genetics 36, S17 - S20 (2004)
http://www.nature.com/ng/journal/v36/n11s/pdf/ng1455.pdf


--S.O.Y. Keita and Rick Kittles. (1997) *The Persistence ofRacial Thinking and the Myth of Racial Divergence. AJPA, 99:3
http://www.councilforresponsiblegenetics.org/pageDocuments/WAURRSZQOE.pdf
---------------- -------

HBD RACE EVOLUTION CLAIMS DEBUNKED BY GENETICISTS
Alan Templeton. "The Genetic and Evolutionary significnce oF Human Races." pp 31-56. IN: J. FiSh (2002) Race and Intelligence: Separating scinnce from myth.
http://img560.imageshack.us/img560/239/templeton1humanracesinf.jpg
http://img685.imageshack.us/img685/2731/templeton2humanracesinf.jpg

IQ claims and miscellaneous data
HBD RACE AND INTELLIGENCE CLAIMS DEBUNKED
 J. FiSh (2002) Race and Intelligence: Separating science from myth.

------------------------------------------





MORE HBD DEBUNKING
-------------------------------- ---------------------



Oubre, A (2011) Race Genes and Ability: Rethinking Ethnic Differences, vol 1 and 2. BTI Press
----------------------------------------------

Krimsky, S, Sloan.K (2011) Race and the Genetic Revolution: Science, Myth, and Culture
-------------------------------


Wicherts and Johnson, 2009. Group differences in the heritability of items and test scores
http://rspb.royalsocietypublishing.org/content/early/2009/04/24/rspb.2009.0238.full


http://nilevalleypeoples.blogspot.com/2013/06/coming-apart-can-murrays-down-with.html


"SELECTION FOR"- "SELECT FOR" HDB CLAIMS DEBUNKED- "SELECTION" IS NOT THE ONLY KEY FORCE DRIVING CHANGE
http://nilevalleypeoples.blogspot.com/2012/12/demic-diffusion-notes-and-tropical.html

http://nilevalleypeoples.blogspot.com/2012/05/7-reasons-libertarians-may-be-wrong.html

http://nilevalleypeoples.blogspot.com/2012/08/cro-magnons-are-us-debunking-claims.html

http://nilevalleypeoples.blogspot.com/2014/03/exploding-nonsense-review-of-cochran_8.html



--Joseph Graves, 2006. What We Know and What We Don’t Know: Human Genetic Variation and the Social Construction of Race
http://raceandgenomics.ssrc.org/Graves/

J. Kahn (2013) How a Drug Becomes "Ethnic" - Yale Journal of Health Policy, Law and Ethics, v4:1
http://digitalcommons.law.yale.edu/cgi/viewcontent.cgi?article=1072&context=yjhple

------------------------------------ -----------------

http://evolution.binghamton.edu/evos/wp-content/uploads/2012/04/PageProofs-Graves_race.pdf

-------------------------------------------- ----------------------------



other links



Evolution, brain size, and the national IQ of peoples ... - Jelte Wicherts 2010
http://wicherts.socsci.uva.nl/wichertsPAIDrejoinder.pdf
------------------------------------

Why national IQs do not support evolutionary theories of intelligence - WIcherts, Borsboom and Dolan 2010
Personality and Individual Differences 48 (2010) 91-96
http://wicherts.socsci.uva.nl/wicherts2010.pdf
----------------------------- -------------

Are intelligence tests measurement invariant over time? by JM Wicherts - ?2004
 --Dolan, Wicherts et al 2004. Investigating the nature of the Flynn effect. Intelligence 32 (2004) 509-537
http://www.iapsych.com/iqmr/fe/LinkedDocuments/wicherts2004.pdf
-------------------------------------------

LYNN AND VANHAVEN'S IQ AND THE WEALTH OF NATIONS DEBUNKED
---------------- -------

www.anth.uconn.edu/faculty/mcbrearty/Pdf/McB%20&%20Brooks%202000%20TRTW.pdf

------------------------

Race and other misadventures: essays in honor of Ashley Montagu... By Larry T. Reynolds, Leonard Lieberman

http://books.google.com/books?id=5DLrgG_MflgC&pg=PA190&dq=r-+k-+selection+races&cd=1#v=onepage&q=r-%20k-%20selection%20races&f=false
--------------------------------

Race and intelligence: separating science from myth. By Jefferson M. Fish. Routledge 2002. See Templeton's detailed article referenced above also inside the book

http://books.google.com/books?id=t9OdPPLIgMAC&pg=PA64&dq=r-+k-+selection+races&cd=7#v=onepage&q=r-%20k-%20selection%20races&f=false
------------------------

http://www.ogiek.org/indepth/what-they-mean.htm
---------------- -------

Oubre, A (2011) Race Genes and Ability: Rethinking Ethnic Differences, vol 1 and 2. BTI Press
For summary see: http://www.skeptic.com/eskeptic/05-02-18/
---------------- -------

http://www.dartmouth.edu/~chance/course/topics/curveball.html

-----------------------------------------------------------

--S OY Keita, R A Kittles, et al. "Conceptualizing human variation," Nature Genetics 36, S17 - S20 (2004)
http://www.nature.com/ng/journal/v36/n11s/pdf/ng1455.pdf


--S.O.Y. Keita and Rick Kittles. (1997) *The Persistence ofRacial Thinking and the Myth of Racial Divergence. AJPA, 99:3
http://www.councilforresponsiblegenetics.org/pageDocuments/WAURRSZQOE.pdf
---------------- -------

HBD RACE EVOLUTION CLAIMS DEBUNKED BY GENETICISTS
Alan Templeton. "The Genetic and Evolutionary significnce oF Human Races." pp 31-56. IN: J. FiSh (2002) Race and Intelligence: Separating scinnce from myth.


HBD RACE AND INTELLIGENCE CLAIMS DEBUNKED
 J. FiSh (2002) Race and Intelligence: Separating science from myth.

------------------------------------------




MORE HBD DEBUNKING
-------------------------------- ---------------------

Oubre, A (2011) Race Genes and Ability: Rethinking Ethnic Differences, vol 1 and 2. BTI Press
----------------------------------------------

Krimsky, S, Sloan.K (2011) Race and the Genetic Revolution: Science, Myth, and Culture
-------------------------------


Wicherts and Johnson, 2009. Group differences in the heritability of items and test scores
http://rspb.royalsocietypublishing.org/content/early/2009/04/24/rspb.2009.0238.full


http://nilevalleypeoples.blogspot.com/2013/06/coming-apart-can-murrays-down-with.html


"SELECTION FOR"- "SELECT FOR" HDB CLAIMS DEBUNKED- "SELECTION" IS NOT THE ONLY KEY FORCE DRIVING CHANGE
http://nilevalleypeoples.blogspot.com/2012/12/demic-diffusion-notes-and-tropical.html

http://nilevalleypeoples.blogspot.com/2012/05/7-reasons-libertarians-may-be-wrong.html

http://nilevalleypeoples.blogspot.com/2012/08/cro-magnons-are-us-debunking-claims.html

http://nilevalleypeoples.blogspot.com/2014/03/exploding-nonsense-review-of-cochran_8.html



--Joseph Graves, 2006. What We Know and What We Don’t Know: Human Genetic Variation and the Social Construction of Race
http://raceandgenomics.ssrc.org/Graves/

J. Kahn (2013) How a Drug Becomes "Ethnic" - Yale Journal of Health Policy, Law and Ethics, v4:1
http://digitalcommons.law.yale.edu/cgi/viewcontent.cgi?article=1072&context=yjhple

------------------------------------ -----------------

http://evolution.binghamton.edu/evos/wp-content/uploads/2012/04/PageProofs-Graves_race.pdf

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