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Saturday, October 18, 2025

Jerry Fodor

From Wikipedia, the free encyclopedia

In 2007
Born
Jerry Alan Fodor

April 22, 1935
New York City, US
DiedNovember 29, 2017 (aged 82)
New York City, US
AwardsJean Nicod Prize (1993)

Education
Alma materColumbia University
Princeton University
ThesisThe Uses of "Use": A Study in the Philosophy of Language (1960)
Doctoral advisorHilary Putnam
Other advisorsSidney Morgenbesser
Philosophical work
Era20th-/21st-century philosophy
RegionWestern philosophy
SchoolAnalytic
InstitutionsRutgers University
Main interestsPhilosophy of mind
Philosophy of language
Cognitive science
Rationalism
Cognitivism
Functionalism
Notable ideasModularity of mind
Language of thought

Jerry Alan Fodor (/ˈfdər/ FOH-dər; April 22, 1935 – November 29, 2017) was an American philosopher and the author of works in the fields of philosophy of mind and cognitive science. His writings in these fields laid the groundwork for the modularity of mind and the language of thought hypotheses, and he is recognized as having had "an enormous influence on virtually every portion of the philosophy of mind literature since 1960." At the time of his death in 2017, he held the position of State of New Jersey Professor of Philosophy, Emeritus, at Rutgers University, and had taught previously at the City University of New York Graduate Center and MIT.

Life and career

Jerry Fodor was born in New York City on April 22, 1935, and was of Jewish descent. He received his degree (summa cum laude) from Columbia University in 1956, where he wrote a senior thesis on Søren Kierkegaard and studied with Sidney Morgenbesser and Arthur Danto. He then earned a PhD in philosophy from Princeton University in 1960, under the direction of Hilary Putnam.

From 1959 to 1986 Fodor was on the faculty of the Massachusetts Institute of Technology. From 1986 to 1988 he was a full professor at the City University of New York (CUNY). From 1988, until his retirement in 2016 as emeritus, he was State of New Jersey Professor of philosophy and cognitive science at Rutgers University.

Besides his interest in philosophy, Fodor followed opera and regularly wrote columns for the London Review of Books on that and other topics. Fodor's first marriage was to the applied psychologist Iris Goldstein, with whom he had one son. After their divorce, he married the linguist Janet Dean. Janet and he lived in Manhattan and had a daughter. He died at home on November 29, 2017.

Philosophical work

Fodor argued in his 1975 book The Language of Thought that mental states, such as beliefs and desires, are relations between individuals and mental representations. He maintained that these representations can only be correctly explained in terms of a language of thought (LOT) in the mind. Furthermore, this language of thought itself is an actually existing thing that is codified in the brain and not just a useful explanatory tool. Fodor adhered to a species of functionalism, maintaining that thinking and other mental processes consist primarily of computations operating on the syntax of the representations that make up the language of thought.

For Fodor, significant parts of the mind, such as perceptual and linguistic processes, are structured in terms of modules, or "organs", which he defines by their causal and functional roles. These modules are relatively independent of each other and of the "central processing" part of the mind, which has a more global and less "domain specific" character. Fodor suggests that the character of these modules permits the possibility of causal relations with external objects. This, in turn, makes it possible for mental states to have contents that are about things in the world. The central processing part, on the other hand, takes care of the logical relations between the various contents and inputs and outputs.

Although Fodor originally rejected the idea that mental states must have a causal, externally determined aspect, in his later years he devoted much of his writing and study to the philosophy of language because of this problem of the meaning and reference of mental contents. His contributions in this area include the so-called asymmetric causal theory of reference and his many arguments against semantic holism. Fodor strongly opposed reductive accounts of the mind. He argued that mental states are multiple realizable and that there is a hierarchy of explanatory levels in science such that the generalizations and laws of a higher-level theory of psychology or linguistics, for example, cannot be captured by the low-level explanations of the behavior of neurons and synapses. He also emerged as a prominent critic of what he characterized as the ill-grounded Darwinian theories of natural selection as an explanation of mind.

Fodor and the nature of mental states

The structure of a mental state, such as that an individual called Mary believes that a man has bitten a dog, as described by Rudolf Carnap, Gottlob Frege, and Fodor. Fodor's scheme says that a person's attitude to something makes use of a mental representation of that thing.

Despite the changes in many of his positions over the years, Fodor's idea that mental states embodying intentionality, propositional attitudes, like beliefs and desires are relational never changed. He attempted to show how mental representations, specifically sentences in the language of thought, are necessary to explain this relational nature of mental states. Fodor considers two alternative hypotheses. The first denies the relational character of mental states, while the second considers mental states as two-place relations. The latter position can be further subdivided into the Carnapian view that such relations are between individuals and sentences of natural languages, and the Fregean view that they are between individuals and the propositions expressed by such sentences. Fodor's own position, instead, is that to properly account for the nature of intentional attitudes, it is necessary to employ a three-place relation between individuals, representations and propositional contents.

Considering mental states as three-place relations in this way, representative realism makes it possible to hold together all of the elements necessary to the solution of this problem. Further, mental representations are not only the objects of beliefs and desires, but are also the domain over which mental processes operate. They can be considered the ideal link between the syntactic notion of mental content and the computational notion of functional architecture. These notions are, according to Fodor, our best explanation of mental processes.

Functional architecture of the mind

Psychological nativism and modularity

The Müller-Lyer illusion, that two identical lines look to be of differing lengths, even if the subject knows they are the same length, persuaded Fodor that mental processes are grouped in discrete modules without access to each other.

Following in the path paved by linguist Noam Chomsky, Fodor developed a strong commitment to the idea of psychological nativism. Nativism postulates the innateness of many cognitive functions and concepts. For Fodor, this emerges naturally out of his criticism of behaviourism and associationism. These criticisms led him to formulate his hypothesis of the modularity of the mind.

Fodor noted the evidence from perceptual errors like the Müller-Lyer illusion that processes such as visually estimating the length of a line are not interfered with by the knowledge that the two lines are actually the same length. He took this to mean that the visual processes were in a separate module from knowledge.

The idea of modularity of mind was presaged by Franz Joseph Gall's 19th century phrenology movement.

The idea can be traced back to the 19th century phrenology movement. Its founder Franz Joseph Gall claimed that mental faculties were associated with specific regions of the brain. Hence, someone's level of intelligence, for example, could be "read off" from the size of a particular bump on his posterior parietal lobe. This simplistic view of modularity has been disproven in the 20th century.

Fodor revived the idea of modularity, without the notion of precise physical localizability, in the 1980s, and became a vocal proponent of it with the 1983 publication of his monograph The Modularity of Mind, where he points to Gall through Bernard Hollander's In search of the soul. Two properties of modularity, informational encapsulation and domain specificity, make it possible to relate functional architecture to mental content. A person's ability to elaborate information independently from their background beliefs allows Fodor to give an atomistic and causal account of mental content. The main idea is that the properties of the contents of mental states can depend, not just on the internal relations of the system of which they are a part, but also on their causal relations with the external world.

Adoption by cognitive and evolutionary psychologists

Fodor's notions of mental modularity, informational encapsulation and domain specificity were taken up and expanded, much to Fodor's chagrin, by cognitive scientists such as Zenon Pylyshyn and evolutionary psychologists such as Steven Pinker and Henry Plotkin. Fodor complained that Pinker, Plotkin and other members of what he sarcastically called "the New Synthesis" had taken modularity and similar ideas way too far. He insisted that the mind is not "massively modular" and that, contrary to these researchers, the mind was a long way from having been explained by the computational, or any other, model. The main reason for this is that most cognition is abductive and global, hence sensitive to all possibly relevant background beliefs to (dis)confirm a belief. This creates the frame problem for the computational theory, because the relevance of a belief is not one of its local, syntactic properties but context-dependent.

Intentional realism

In A Theory of Content and Other Essays (1990), Fodor takes up another of his central notions: the question of the reality of mental representations. He sought to justify representational realism, so as to justify the idea that the contents of mental states are expressed in symbolic structures such as those of the LOT.

Fodor's criticism of Dennett

Fodor starts with some criticisms of so-called standard realism. This view is characterized, according to Fodor, by two distinct assertions. One of these regards the internal structure of mental states and asserts that such states are non-relational. The other concerns the semantic theory of mental content and asserts that there is an isomorphism between the causal roles of such contents and the inferential web of beliefs. Among modern philosophers of mind, the majority view seems to be that the first of these two assertions is false, but that the second is true. Fodor departs from this view in accepting the truth of the first thesis but rejecting strongly the truth of the second.

In particular, Fodor criticizes the instrumentalism of Daniel Dennett. Dennett maintains that it is possible to be realist with regard to intentional states without having to commit oneself to the reality of mental representations. Now, according to Fodor, if one remains at this level of analysis, then there is no possibility of explaining why the intentional strategy works:

There is ... a standard objection to instrumentalism ...: it is difficult to explain why the psychology of beliefs/desires works so well, if the psychology of beliefs/desires is, in fact, false.... As Putnam, Boyd and others have emphasized, from the predictive successes of a theory to the truth of that theory there is surely a presumed inference; and this is even more likely when ... we are dealing with the only theory in play which is predictively crowned with success. It is not obvious ... why such a presumption should not militate in favour of a realist conception ... of the interpretations of beliefs/desires.

Productivity, systematicity and thought

Building on Chomsky

Fodor also has positive arguments in favour of the reality of mental representations in terms of the LOT. He maintains that if language is the expression of thoughts and language is systematic, then thoughts must also be systematic. Fodor draws on the work of Noam Chomsky to both model his theory of the mind and to refute alternative architectures such as connectionism. Systematicity in natural languages was explained by Chomsky in terms of two more basic concepts: productivity and compositionality.

Productivity is a representational system's unbounded ability to generate new representations from a given set of symbols, using its cognitive architecture. "John", "loves", and "Mary" allow for the construction of the sentences "John loves Mary" and "Mary loves John". Fodor's language of thought theorizes that representations are decomposable into constituent parts, and these decomposed representations are built into new strings.

More important than productivity is systematicity since it does not rely on questionable idealizations about human cognition. The argument states that a cognizer is able to understand some sentence in virtue of understanding another. For example, no one who understands "John loves Mary" is unable to understand "Mary loves John", and no one who understands "P and Q" is unable to understand "P". Systematicity itself is rarely challenged as a property of natural languages and logics, but some challenge that thought is systematic in the same way languages are. Still others from the connectionist tradition have tried to build non-classical networks that can account for the apparent systematicity of language.

The fact that systematicity and productivity depend on the compositional structure of language means that language has a combinatorial semantics. If thought also has such a combinatorial semantics, then, Fodor stated, there must be a language of thought.

Formalizing thought processes

The second argument that Fodor provides in favour of representational realism involves the processes of thought. This argument touches on the relation between the representational theory of mind and models of its architecture. If the sentences of Mentalese require unique processes of elaboration then they require a computational mechanism of a certain type. The syntactic notion of mental representations goes hand in hand with the idea that mental processes are calculations which act only on the form of the symbols which they elaborate. And this is the computational theory of the mind. Consequently, the defence of a model of architecture based on classic artificial intelligence passes inevitably through a defence of the reality of mental representations.

For Fodor, this formal notion of thought processes also has the advantage of highlighting the parallels between the causal role of symbols and the contents which they express. In his view, syntax plays the role of mediation between the causal role of the symbols and their contents. The semantic relations between symbols can be "imitated" by their syntactic relations. The inferential relations which connect the contents of two symbols can be imitated by the formal syntax rules which regulate the derivation of one symbol from another.

The nature of content

From the beginning of the 1980s, Fodor adhered to a causal notion of mental content and of meaning. This idea of content contrasts sharply with the inferential role semantics to which he subscribed earlier in his career. Fodor went on to criticize inferential role semantics (IRS) because its commitment to an extreme form of holism excludes the possibility of a true naturalization of the mental. But naturalization must include an explanation of content in atomistic and causal terms.

Anti-holism

Fodor, while following W.V.O. Quine's confirmation holism, criticised semantic holism, the idea that every connection of a concept is part of its meaning. The identity of the content of a mental state, under semantic holism, can only be determined by the totality of its epistemic bonds. Fodor argued that this makes the realism of mental states an impossibility:

If people differ in an absolutely general way in their estimations of epistemic relevance, and if we follow the holism of meaning and individuate intentional states by way of the totality of their epistemic bonds, the consequence will be that two people (or, for that matter, two temporal sections of the same person) will never be in the same intentional state. Therefore, two people can never be subsumed under the same intentional generalizations. And, therefore, intentional generalization can never be successful. And, therefore again, there is no hope for an intentional psychology.

The asymmetric causal theory

Having criticized the idea that semantic evaluation concerns only the internal relations between the units of a symbolic system, Fodor can adopt an externalist position with respect to mental content and meaning. For Fodor, later in his life, the problem of naturalization of the mental is tied to the possibility of giving "the sufficient conditions for which a piece of the world is relative to (expresses, represents, is true of) another piece" in non-intentional and non-semantic terms. If this goal is to be achieved within a representational theory of the mind, then the challenge is to devise a causal theory which can establish the interpretation of the primitive non-logical symbols of the LOT. Fodor's initial proposal is that what determines that the symbol for "water" in Mentalese expresses the property H2O is that the occurrences of that symbol are in certain causal relations with water. The intuitive version of this causal theory is what Fodor calls the "Crude Causal Theory". According to this theory, the occurrences of symbols express the properties which are the causes of their occurrence. The term "horse", for example, says of a horse that it is a horse. In order to do this, it is necessary and sufficient that certain properties of an occurrence of the symbol "horse" be in a law-like relation with certain properties which determine that something is an occurrence of horse.

The main problem with this theory is that of erroneous representations. There are two unavoidable problems with the idea that "a symbol expresses a property if it is ... necessary that all and only the presences of such a property cause the occurrences". The first is that not all horses cause occurrences of horse. The second is that not only horses cause occurrences of horse. Sometimes the A(horses) are caused by A (horses), but at other times—when, for example, because of the distance or conditions of low visibility, one has confused a cow for a horse—the A (horses) are caused by B (cows). In this case the symbol A doesn't express just the property A, but the disjunction of properties A or B. The crude causal theory is therefore incapable of distinguishing the case in which the content of a symbol is disjunctive from the case in which it isn't. This gives rise to what Fodor calls the "problem of disjunction".

Fodor responds to this problem with what he defines as "a slightly less crude causal theory". According to this approach, it is necessary to break the symmetry at the base of the crude causal theory. Fodor must find some criterion for distinguishing the occurrences of A caused by As (true) from those caused by Bs (false). The point of departure, according to Fodor, is that while the false cases are ontologically dependent on the true cases, the reverse is not true. There is an asymmetry of dependence, in other words, between the true contents (A= A) and the false ones (A=A or B). The first can subsist independently of the second, but the second can occur only because of the existence of the first:

From the point of view of semantics, errors must be accidents: if in the extension of "horse" there are no cows, then it cannot be required for the meaning of "horse" that cows be called horses. On the other hand, if "horse" did not mean that which it means, and if it were an error for horses, it would never be possible for a cow to be called "horse". Putting the two things together, it can be seen that the possibility of falsely saying "this is a horse" presupposes the existence of a semantic basis for saying it truly, but not vice versa. If we put this in terms of the crude causal theory, the fact that cows cause one to say "horse" depends on the fact that horses cause one to say "horse"; but the fact that horses cause one to say "horse" does not depend on the fact that cows cause one to say "horse"...

Functionalism

During the 1960s, philosophers such as Donald Davidson, Hilary Putnam, and Fodor tried to resolve the puzzle of developing a way to preserve the explanatory efficacy of mental causation and folk psychology while adhering to a materialist vision of the world which did not violate the "generality of physics". Their proposal was, first of all, to reject the then-dominant theories in philosophy of mind: behaviorism and the type identity theory. The problem with logical behaviorism was that it failed to account for causation between mental states and such causation seems to be essential to psychological explanation, especially if one considers that behavior is not an effect of a single mental event/cause but is rather the effect of a chain of mental events/causes. The type-identity theory, on the other hand, failed to explain the fact that radically different physical systems can find themselves in the identical mental state. Besides being deeply anthropocentric (why should humans be the only thinking organisms in the universe?), the identity-type theory also failed to deal with accumulating evidence in the neurosciences that every single human brain is different from all the others. Hence, the impossibility of referring to common mental states in different physical systems manifests itself not only between different species but also between organisms of the same species.

An illustration of multiple realizability. M stands for mental and P stand for physical. The diagram shows that more than one P can instantiate one M, but not vice versa. Causal relations between states are represented by the arrows (M1 goes to M2, etc.).

One can solve these problems, according to Fodor, with functionalism, a hypothesis which was designed to overcome the failings of both dualism and reductionism. What is important is the function of a mental state regardless of the physical substrate which implements it. The foundation for this view lies in the principle of the multiple realizability of the mental. Under this view, for example, I and a computer can both instantiate ("realize") the same functional state though we are made of completely different material stuff (see graphic at right). On this basis functionalism can be classified as a form of token materialism.

Evolution

Fodor and the biolinguist Massimo Piattelli-Palmarini co-authored the book What Darwin Got Wrong (2010). In the same, the authors argue that much neo-Darwinian literature is "distressingly uncritical" and that Charles Darwin's theory of evolution "overestimates the contribution the environment makes in shaping the phenotype of a species and correspondingly underestimates the effects of endogenous variables". Evolutionary biologist Jerry Coyne describes this book as "a profoundly misguided critique of natural selection" and "as biologically uninformed as it is strident". Moral philosopher and anti-scientism author Mary Midgley praises What Darwin Got Wrong as "an overdue and valuable onslaught on neo-Darwinist simplicities".

Awards and honors

Fodor was a member of the American Academy of Arts and Sciences. He received numerous awards and honors: New York State Regent's Fellowship, Woodrow Wilson Fellowship (Princeton University), Chancellor Greene Fellow (Princeton University), Fulbright Fellowship (University of Oxford), Fellow at the Center for Advanced Study in the Behavioral Sciences, and a Guggenheim Fellowship. He won the first Jean Nicod Prize for philosophy of mind and cognitive philosophy in 1993. His lecture series for the Prize was published as The Elm and the Expert: Mentalese and Its Semantics (1995). In 1996–1997, Fodor delivered the John Locke Lectures at the University of Oxford, titled Concepts: Where Cognitive Science Went Wrong, which went on to become a 1998 book. He also delivered the Patrick Romanell Lecture on Philosophical Naturalism (2004) and the Royce Lecture on Philosophy of Mind (2002) to the American Philosophical Association, of whose Eastern Division he served as President (2005–2006). In 2005, he won the Mind & Brain Prize.

Criticism

Philosophers of diverse orientations have challenged many of Fodor's ideas.

Simon Blackburn

Simon Blackburn in Spreading the Word (1984) has accused the language of thought hypothesis of falling prey to an infinite regress. If a person understands a word like "dog" through a mental representation which indicates that the word denotes dogs, then a regress takes place. Fodor replies that the language of thought does not involve denoting things. On this view, the process of reasoning works directly to give a result but does not allow the person to examine how it works.

Daniel Dennett

In 1981, Daniel Dennett formulated another argument against the LOT. Dennett suggested that it would seem, on the basis of the evidence of our behavior toward computers but also with regard to some of our own unconscious behavior, that explicit representation is not necessary for the explanation of propositional attitudes. During a game of chess with a computer program, we often attribute such attitudes to the computer, saying such things as "It thinks that the queen should be moved to the left." We attribute propositional attitudes to the computer and this helps us to explain and predict its behavior in various contexts. Yet no one would suggest that the computer is actually thinking or believing somewhere inside its circuits the equivalent of the propositional attitude "I believe I can kick this guy's butt" in Mentalese. The same is obviously true, suggests Dennett, of many of our everyday automatic behaviors such as "desiring to breathe clear air" in a stuffy environment.

Kent Bach

Linguists and philosophers of language such as Kent Bach have criticized Fodor's self-proclaimed "extreme" concept nativism. Bach takes Fodor to task for his criticisms of lexical semantics and polysemy. Fodor claims that there is no lexical structure to such verbs as "keep", "get", "make" and "put". He suggests that, alternatively, "keep" simply expresses the concept KEEP (Fodor capitalizes concepts to distinguish them from properties, names or other such entities). If there is a straightforward one-to-one mapping between individual words and concepts, "keep your clothes on", "keep your receipt" and "keep washing your hands" will all share the same concept of KEEP under Fodor's theory. This concept presumably locks on to the unique external property of keeping. But, if this is true, then RETAIN must pick out a different property in RETAIN YOUR RECEIPT, since one can't retain one's clothes on or retain washing one's hands. Fodor's theory also has a problem explaining how the concept FAST contributes, differently, to the contents of FAST CAR, FAST DRIVER, FAST TRACK, and FAST TIME. Whether or not the differing interpretations of "fast" in these sentences are specified in the semantics of English, or are the result of pragmatic inference, is a matter of debate. Fodor's own response to this kind of criticism is expressed bluntly in Concepts: "People sometimes used to say that exist must be ambiguous because look at the difference between 'chairs exist' and 'numbers exist'. A familiar reply goes: the difference between the existence of chairs and the existence of numbers seems, on reflection, strikingly like the difference between numbers and chairs. Since you have the latter to explain the former, you don't also need 'exist' to be polysemic."

Some critics find it difficult to accept Fodor's insistence that a large, perhaps implausible, number of concepts are primitive and undefinable. For example, Fodor considers such concepts as EFFECT, ISLAND, TRAPEZOID, and WEEK to be all primitive, innate and unanalyzable because they all fall into the category of what he calls "lexical concepts" (those for which our language has a single word). Against this view, Bach argues that the concept VIXEN is almost certainly composed out of the concepts FEMALE and FOX, BACHELOR out of SINGLE and MALE, and so on.

Fiona Cowie

In her 1999 book What's Within, Fiona Cowie argued against Fodor's innatist view, preferring a John Locke-style empiricism. Fodor replied at length in a 1999 article titled "Doing Without What's Within; Fiona Cowie's Critique of Nativism", stating that he was aiming at a position halfway between nativism and empiricism.

Sociolinguistics

From Wikipedia, the free encyclopedia
https://en.wikipedia.org/wiki/Sociolinguistics

Sociolinguistics is the descriptive, scientific study of how language is shaped by, and used differently within, any given society. The field largely looks at how a language varies between distinct social groups and under the influence of assorted cultural norms, expectations, and contexts, including how that variation plays a role in language change. Sociolinguistics combines the older field of dialectology with the social sciences in order to identify regional dialects, sociolects, ethnolects, and other sub-varieties and styles within a language.

A major branch of linguistics since the second half of the 20th century, sociolinguistics is closely related to and can partly overlap with pragmatics, linguistic anthropology, and sociology of language, the latter focusing on the effect of language back on society. Sociolinguistics' historical interrelation with anthropology can be observed in studies of how language varieties differ between groups separated by social variables (e.g., ethnicity, religion, status, gender, level of education, age, etc.) or geographical barriers (a mountain range, a desert, a river, etc.). Such studies also examine how such differences in usage and in beliefs about usage produce and reflect social or socioeconomic classes. As the usage of a language varies from place to place, language usage also varies among social classes, and some sociolinguists study these sociolects.

Studies in the field of sociolinguistics use a variety of research methods including ethnography and participant observation, analysis of audio or video recordings of real life encounters or interviews with members of a population of interest. Some sociolinguists assess the realization of social and linguistic variables in the resulting speech corpus. Other research methods in sociolinguistics include matched-guise tests (in which listeners share their evaluations of linguistic features they hear), dialect surveys, and analysis of preexisting corpora.

Sociolinguistics in history

Beginnings

The social aspects of language were in the modern sense first studied by Indian and Japanese linguists in the 1930s, and also by forerunners in Denmark and Switzerland around the turn of the 20th century, but none received much attention in the West until much later. The study of the social motivation of language change, on the other hand, has its foundation in the wave model of the late 19th century. The first attested use of the term sociolinguistics was by Thomas Callan Hodson in the title of his 1939 article "Sociolinguistics in India" published in Man in India.

Dialectology is an old field, and in the early 20th century, dialectologists such as Hans Kurath and Raven I. McDavid Jr. initiated large scale surveys of dialect regions in the U.S.

Western contributions

The study of sociolinguistics in the West was pioneered by linguists such as Charles A. Ferguson or William Labov in the US and Basil Bernstein in the UK. In the 1960s, William Stewart and Heinz Kloss introduced the basic concepts for the sociolinguistic theory of pluricentric languages, which describes how standard language varieties differ between nations, e.g. regional varieties of English versus pluricentric "English"regional standards of German versus pluricentric "German"Bosnian, Croatian, Montenegrin, and Serbian versus pluricentric "Serbo-Croatian". Dell Hymes, one of the founders of linguistic anthropology, is credited with developing an ethnography-based sociolinguistics and is the founder of the journal Language in Society. His focus on ethnography and communicative competence contributed to his development of the SPEAKING method: an acronym for setting, participants, ends, act sequence, keys, instrumentalities, norms, and genres that is widely recognized as a tool to analyze speech events in their cultural context.

Applications

Sociolinguistics can be divided into subfields, which make use of different research methods, and have different goals. Dialectologists survey people through interviews, and compile maps. Ethnographers such as Dell Hymes and his students often live amongst the people they are studying. Conversation analysts such as Harvey Sacks and interactional sociolinguists such as John J. Gumperz record audio or video of natural encounters, and then analyze the tapes in detail. Sociolinguists tend to be aware of how the act of interviewing might affect the answers given.

Some sociolinguists study language on a national level among large populations to find out how language is used as a social institution. William Labov, a Harvard and Columbia University graduate, is often regarded as the founder of variationist sociolinguistics which focuses on the quantitative analysis of variation and change within languages, making sociolinguistics a scientific discipline.

For example, a sociolinguistics-based translation framework states that a linguistically appropriate translation cannot be wholly sufficient to achieve the communicative effect of the source language; the translation must also incorporate the social practices and cultural norms of the target language. To reveal social practices and cultural norms beyond lexical and syntactic levels, the framework includes empirical testing of the translation using methods such as cognitive interviewing with a sample population.

A commonly studied source of variation is regional dialects. Dialectology studies variations in language based primarily on geographic distribution and their associated features. Sociolinguists concerned with grammatical and phonological features that correspond to regional areas are often called dialectologists.

Sociolinguistic interview

The sociolinguistic interview is the foundational method of collecting data for sociolinguistic studies, allowing the researcher to collect large amounts of speech from speakers of the language or dialect being studied. The interview takes the form of a long, loosely structured conversation between the researcher and the interview subject; the researcher's primary goal is to elicit the vernacular style of speech: the register associated with everyday casual conversation. This goal is complicated by the observer's paradox: the researcher is trying to elicit the style of speech that would be used if the interviewer were not present.

To that end, a variety of techniques may be used to reduce the subject's attention to the formality and artificiality of the interview setting. For example, the researcher may attempt to elicit narratives of memorable events from the subject's life, such as fights or near-death experiences; the subject's emotional involvement in telling the story is thought to distract their attention from the formality of the context. Some researchers interview multiple subjects together to allow them to converse more casually with one other than they would with the interviewer alone. The researcher may then study the effects of style-shifting on language by comparing a subject's speech style in more vernacular contexts, such as narratives of personal experience or conversation between subjects, with the more careful style produced when the subject is more attentive to the formal interview setting. The correlations of demographic features such as age, gender, and ethnicity with speech behavior may be studied by comparing the speech of different interview subjects.

Fundamental concepts

While the study of sociolinguistics is very broad, there are a few fundamental concepts on which many sociolinguistic inquiries depend.

Speech community

Speech community is a concept in sociolinguistics that describes a distinct group of people who use language in a unique and mutually accepted way among themselves. This is sometimes referred to as a Sprechbund.

To be considered part of a speech community, one must have a communicative competence. That is, the speaker has the ability to use language in a way that is appropriate in the given situation. It is possible for a speaker to be communicatively competent in more than one language.

Demographic characteristics such as areas or locations have helped to create speech community boundaries in speech community concept. Those characteristics can assist exact descriptions of specific groups' communication patterns.

Speech communities can be members of a profession with a specialized jargon, distinct social groups like high school students or hip hop fans, or even tight-knit groups like families and friends. Members of speech communities will often develop slang or specialized jargon to serve the group's special purposes and priorities. This is evident in the use of lingo within sports teams.

Community of Practice allows for sociolinguistics to examine the relationship between socialization, competence, and identity. Since identity is a very complex structure, studying language socialization is a means to examine the micro-interactional level of practical activity (everyday activities). The learning of a language is greatly influenced by family, but it is supported by the larger local surroundings, such as school, sports teams, or religion. Speech communities may exist within a larger community of practice.

High-prestige and low-prestige varieties

Crucial to sociolinguistic analysis is the concept of prestige; certain speech habits are assigned a positive or a negative value, which is then applied to the speaker. This can operate on many levels. It can be realized on the level of the individual sound/phoneme, as Labov discovered in investigating pronunciation of the post-vocalic /r/ in the Northeastern United States, or on the macro scale of language choice, as is realized in the various diglossia that exist throughout the world, with the one between Swiss German and High German being perhaps most well known. An important implication of the sociolinguistic theory is that speakers 'choose' a variety when making a speech act, whether consciously or subconsciously.

The terms acrolectal (high) and basilectal (low) are also used to distinguish between a more standard dialect and a dialect of less prestige.

It is generally assumed that non-standard language is low-prestige language. However, in certain groups, such as traditional working-class neighborhoods, standard language may be considered undesirable in many contexts because the working-class dialect is generally considered a powerful in-group marker. Historically, humans tend to favor those who look and sound like them, and the use of nonstandard varieties (even exaggeratedly so) expresses neighborhood pride and group and class solidarity. The desirable social value associated with the use of non-standard language is known as covert prestige. There will thus be a considerable difference in use of non-standard varieties when going to the pub or having a neighborhood barbecue compared to going to the bank. One is a relaxed setting, likely with familiar people, and the other has a business aspect to it in which one feels the need to be more professional.

Social network

Understanding language in society means that one also has to understand the social networks in which language is embedded. A social network is another way of describing a particular speech community in terms of relations between individual members in a community. A network could be loose or tight depending on how members interact with each other. For instance, an office or factory may be considered a tight community because all members interact with each other. A large course with 100+ students would be a looser community because students may only interact with the instructor and maybe 1–2 other students. A multiplex community is one in which members have multiple relationships with each other. For instance, in some neighborhoods, members may live on the same street, work for the same employer and even intermarry.

The looseness or tightness of a social network may affect speech patterns adopted by a speaker. For instance, Sylvie Dubois and Barbara Horvath found that speakers in one Cajun Louisiana community were more likely to pronounce English "th" [θ] as [t] (or [ð] as [d]) if they participated in a relatively dense social network (i.e. had strong local ties and interacted with many other speakers in the community), and less likely if their networks were looser (i.e. fewer local ties).

A social network may apply to the macro level of a country or a city, but also to the interpersonal level of neighborhoods or a single family. Recently, social networks have been formed by the Internet through online chat rooms, Facebook groups, organizations, and online dating services.

Differences according to class

Sociolinguistics as a field distinct from dialectology was pioneered through the study of language variation in urban areas. Whereas dialectology studies the geographic distribution of language variation, sociolinguistics focuses on other sources of variation, among them class. Class and occupation are among the most important linguistic markers found in society. One of the fundamental findings of sociolinguistics, which has been hard to disprove, is that class and language variety are related. Members of the working class tend to speak less of what is deemed standard language, while the lower, middle, and upper middle class will, in turn, speak closer to the standard. However, the upper class, even members of the upper middle class, may often speak 'less' standard than the middle class. This is because not only class but class aspirations, are important. One may speak differently or cover up an undesirable accent to appear to have a different social status and fit in better with either those around them, or how they wish to be perceived.

Class aspiration

Studies, such as those by William Labov in the 1960s, have shown that social aspirations influence speech patterns. This is also true of class aspirations. In the process of wishing to be associated with a certain class (usually the upper class and upper middle class) people who are moving in that direction socio-economically may adjust their speech patterns to sound like them. However, not being native upper-class speakers, they often hypercorrect, which involves overcorrecting their speech to the point of introducing new errors. The same is true for individuals moving down in socio-economic status.

In any contact situation, there is a power dynamic, be it a teacher-student or employee-customer situation. This power dynamic results in a hierarchical differentiation between languages.

Non-standard dialect
(associated with lower classes)
Standard dialect
(associated with higher classes)
It looks like it ain't gonna rain today. It looks as if it isn't going to rain today.
You give it to me yesterday. You gave it to me yesterday.
Y'gotta do it the right way. You have to do it the right way.

Social language codes

Basil Bernstein, a well-known British sociolinguist, devised in his book, Elaborated and restricted codes: their social origins and some consequences, a method for categorizing language codes according to variable emphases on verbal and extraverbal communication. He claimed that factors like family orientation, social control, verbal feedback, and possibly social class contributed to the development of the two codes: elaborated and restricted.

Restricted code

According to Basil Bernstein, the restricted code exemplified the predominance of extraverbal communication, with an emphasis on interpersonal connection over individual expression. His theory places the code within environments that operate according to established social structures that predetermine the roles of their members in which the commonality of interests and intents from a shared local identity creates a predictability of discrete intent and therefore a simplification of verbal utterances. Such environments may include military, religious, and legal atmospheres; criminal and prison subcultures; long-term married relationships; and friendships between children.

The strong bonds between speakers often renders explicit verbal communication unnecessary and individual expression irrelevant. However, simplification is not a sign of a lack of intelligence or complexity within the code; rather, communication is performed more through extraverbal means (facial expression, touch, etc.) in order to affirm the speakers' bond. Bernstein notes the example of a young man asking a stranger to dance since there is an established manner of asking, yet communication is performed through physical graces and the exchange of glances.

As such, implied meaning plays a greater role in this code than in the elaborated code. Restricted code also operates to unify speakers and foster solidarity.

Elaborated code

Basil Bernstein defined 'elaborated code' according to its emphasis on verbal communication over extraverbal. This code is typical in environments where a variety of social roles are available to the individual, to be chosen based upon disposition and temperament. Most of the time, speakers of elaborated code use a broader lexicon and demonstrate less syntactic predictability than speakers of restricted code. The lack of predetermined structure and solidarity requires explicit verbal communication of discrete intent by the individual to achieve educational and career success.

Bernstein notes with caution the association of the code with upper classes (while restricted code is associated with lower classes) since the abundance of available resources allows persons to choose their social roles. He warns, however, that studies associating the codes with separate social classes used small samples and were subject to significant variation.

He also asserts that elaborated code originates from differences in social context, rather than intellectual advantages. As such, elaborated code differs from restricted code according to the context-based emphasis on individual advancement over assertion of social/community ties.

The codes and child development

Bernstein explains language development according to the two codes in light of their fundamentally different values. For instance, a child exposed solely to restricted code learns extraverbal communication over verbal, and therefore may have a less extensive vocabulary than a child raised with exposure to both codes. While there is no inherent lack of value to restricted code, a child without exposure to elaborated code may encounter difficulties upon entering formal education, in which standard, clear verbal communication and comprehension is necessary for learning and effective interaction both with instructors and other students from differing backgrounds. As such, it may be beneficial for children who have been exposed solely to restricted code to enter pre-school training in elaborated code in order to acquire a manner of speaking that is considered appropriate and widely comprehensible within the education environment.

Additionally, Bernstein notes several studies in language development according to social class. In 1963, the Committee for Higher Education conducted a study on verbal IQ that showed a deterioration in individuals from lower working classes ages 8–11 and 11–15 years in comparison to those from middle classes (having been exposed to both restricted and elaborated codes). Additionally, studies by Bernstein, Venables, and Ravenette, as well as a 1958 Education Council report, show a relative lack of success on verbal tasks in comparison to extraverbal in children from lower working classes (having been exposed solely to restricted code).

Contradictions

The idea of these social language codes from Bernstein contrast with famous linguist Noam Chomsky's ideas. Chomsky, deemed the "father of modern linguistics", argues that there is a universal grammar, meaning that humans are born with an innate capacity for linguistic skills like sentence-building. This theory has been criticized by several scholars of linguistic backgrounds because of the lack of proven evolutionary feasibility and the fact that different languages do not have universal characteristics.

Sociolinguistic variation

The study of language variation is concerned with social constraints determining language in its contextual environment. The variations will determine some of the aspects of language like the sound, grammar, and tone in which people speak, and even non-verbal cues. Code-switching is the term given to the use of different varieties of language depending on the social situation. This is commonly used among the African-American population in the United States. There are several different types of age-based variation one may see within a population as well such as age range, age-graded variation, and indications of linguistic change in progress. The use of slang can be a variation based on age. Younger people are more likely to recognize and use today's slang while older generations may not recognize new slang, but might use slang from when they were younger.

Variation may also be associated with gender, as men and women, on average, tend to use slightly different language styles. These differences are typically quantitative rather than qualitative. In other words, while women may use certain speaking styles more frequently than men, the distinction is comparable to height differences between the sexes—on average, men are taller than women, yet some women are taller than some men. Similar variations in speech patterns include differences in pitch, tone, speech fillers, interruptions, and the use of euphemisms, etc.

These gender-based differences in communication extend beyond face-to-face interactions and are also evident in digital spaces. Despite the continuous evolution of social media platforms, cultural and societal norms continue to shape online interactions. For instance, men and women often adopt different non-verbal cues and roles in virtual conversations. However, when it comes to fundamental aspects of communication—such as spoken language, active listening, providing feedback, understanding context, selecting communication methods, and managing conflicts—their approaches tend to be more similar than different.

Beyond these stylistic differences, research suggests that gendered language patterns are also influenced by social expectations and power dynamics. Women, for instance, are more likely to use hedging expressions (e.g., "I think" or "perhaps") and tag questions ("isn't it?") to soften their statements and promote conversational cooperation. Meanwhile, men tend to adopt more assertive and direct speech patterns, reflecting broader societal norms that associate masculinity with dominance and authority.

Variation in language can also come from ethnicity, economic status, level of education, etc.

Friday, October 17, 2025

Genetic history of Africa

From Wikipedia, the free encyclopedia
Population structure of African populations in a broad context. ADMIXTURE analysis of 2,194 individuals from 81 populations for 16,420 SNPs reveals both well-established and novel ancestry components in African populations.

The genetic history of Africa summarizes the genetic makeup and population history of African populations in Africa, composed of the overall genetic history, including the regional genetic histories of North Africa, West Africa, East Africa, Central Africa, and Southern Africa, as well as the recent origin of modern humans in Africa. The Sahara served as a trans-regional passageway and place of dwelling for people in Africa during various humid phases and periods throughout the history of Africa. It also served as a biological barrier that restricted geneflow between the northern and central parts of Africa since its desertification, contributing to the diverse and distinct population structures on the continent. Nonetheless, this did not stop contact between peoples north and south of the Sahara at various points, especially in prehistoric times when the climate conditions were warmer and wetter.

Overview

(A) the origin of the 46 African ethnic groups used in the analysis; ethnic groups from similar regions are given the same colour, but different shapes. (B) PCA shows that the first major axis of variation in Africa (PC1, y-axis) splits southern groups from the rest of Africa, each symbol represents an individual; PC2 (x-axis) reflects ethno-linguistic differences, with Niger-Congo and Nilo-Saharan speakers split from Afroasiatic speakers. (C) The third principal component (PC3, x-axis) represents geographical separation of Niger-Congo speakers, forming a cline from west to east Africans.

The peoples of Africa are characterized by regional genetic substructure and heterogeneity, depending on the respective ethno-linguistic identity, and, in part, explainable by the "multiregional evolution" of modern human lineages in various multiple regions of the African continent, as well as later admixture events, including back-migrations from Eurasia, of both highly differentiated West and East Eurasian components.

Africans' genetic ancestry is largely partitioned by geography and language family, with populations belonging to the same ethno-linguistic groupings showing high genetic homogeneity and coherence. Gene flow, consistent with both short- and long-range migration events followed by extensive admixture and bottleneck events, have influenced the regional genetic makeup and demographic structure of Africans. The historical Bantu expansion had lasting impacts on the modern demographic make up of Africa, resulting in a greater genetic and linguistic homogenization. Genetic, archeologic, and linguistic studies added extra insight into this movement: "Our results reveal a genetic continuum of Niger–Congo speaker populations across the continent and extend our current understanding of the routes, timing and extent of the Bantu migration."

Overall, different African populations display genetic diversity and substructure, but can be clustered in distinct but partially overlapping groupings:

  • Khoisan or 'South African hunter-gatherers' from Southern Africa represented by the Khoisan peoples; they are associated with the deepest divergence (c. 270,000 years ago) of human genetic diversity, forming a distinct cluster of their own. They subsequently diverged into a Northern and Southern subgroup, c. 30,000 years ago.
  • 'Central African hunter-gatherers' or 'Rain forest hunter-gatherers' (Pygmies) of Central Africa, represented by the Biaka and Mbuti; associated with another deep divergence (c. 220,000 years ago). They subsequently diverged into an Eastern and Western subgroup, c. 20,000 years ago.
  • "Ancestral Eurasians" represent the ancestral population of modern Eurasians shortly before the Out-of-Africa expansion; they are inferred to have diverged from other African populations, most likely somewhere in Northeast Africa, c. 70,000 years ago.
  • The various Afroasiatic-speakers from Northern Africa and the Horn of Africa, are suggested to have diverged from other African groups c. 50,000 years ago, but currently insufficient data and geneflow from other groups complicate an accurate estimation of the divergence date. Afroasiatic-speaking populations also display variable amounts of West Asian (primarily Natufian-like, but also Neolithic Anatolian and Iranian) admixtures from Eurasian backflow movements, with the remainder being primarily from autochthonous African genetic clusters, associated with Nilotic-like ancestry. They also display affinity for the Paleolithic North African Taforalt specimens of the Iberomaurusian culture.
  • 'Eastern African hunter-gatherers', represented by Hadza, Sandawe, Omotic-speakers, and the ancient Mota specimen; their phylogenetic relationship to other populations is not clear, but they display affinity to modern East and West African populations, and harbor Khoesan-like geneflow along a Northeast to Southwest cline, as well as later (West) Eurasian admixtures, but at lower amounts than among Afroasiatic-speakers.
  • "Ancient East Africans" or "Ancestral West/East Africans" associated with the common ancestor of modern Niger-Congo and Nilo-Saharan-speakers originated around 28,000 years ago, likely in the Nile Valley region. They subsequently diverged at c. 18,000 years ago into the ancestors of West and West-Central African Niger-Congo and Bantu-speakers, and into the East African Nilo-Saharan/Nilotic-speakers. They represent the dominant and most widespreaded ancestry component of modern Africa, and are associated with relative recent population expansions linked to agriculture and pastoralist lifestyles. Genetic data indicates affinity for older hunter-gatherer groups in East Africa, but their exact relationship remains unclear. There is evidence for limited geneflow (9–13%) from a human ghost lineage, referred to as 'West African foragers' with a deeper or equally deep divergence time than 'Khoisan hunter-gatherers', into modern West Africans.
  • Austronesian-speaking Malagasy people in Madagascar have received significant East/Southeast Asian admixture associated with the Austronesian expansion, with the remainder ancestry being primarily associated with West-Central and East African components. The estimated date of geneflow between these sources is c. 2,200 years ago.
Geographic location of the samples analyzed in this study (A). PCA of the Khoe-San individuals, Eurasians, West and East Africans before (unmasked, B) and after (masked, C) applying the local ancestry pipeline (146,696 independent SNPs).

Indigenous Africans

The term 'indigenous Africans' refers to the populations with primarily indigenous (non-Eurasian) ancestries, consisting of Niger–Congo speakers, Nilo-Saharan speakers, the divergent and diverse Khoisan grouping, as well as of several unclassified or isolated ethnolinguistic groupings (see unclassified languages of Africa). The origin of the Afroasiatic languages remains disputed, with some proposing a Middle Eastern origin, while others support an African origin with varying degrees of Eurasian and African components. The Austronesian languages originated in southern East Asia, and later expanded outgoing from the Philippines.

PCA plot of genetic variation of worldwide populations. (A) Geographic coordinates of 53 populations. (B) Procrustes-transformed PCA plot of genetic variation.

The Niger–Congo languages probably originated in or near the area where these languages were spoken prior to Bantu expansion (i.e. West Africa or Central Africa). Its expansion may have been associated with the expansion of agriculture, in the African Neolithic period, following the desiccation of the Sahara in c. 3500 BCE. Proto-Niger-Congo may have originated about 10,000 years before present in the "Green Sahara" of Africa (roughly the Sahel and southern Sahara), and that its dispersal can be correlated with the spread of the bow and arrow by migrating hunter-gatherers, which later developed agriculture.

Although the validity of the Nilo-Saharan family remains controversial, the region between Chad, Sudan, and the Central African Republic is seen as a likely candidate for its homeland prior to its dispersal around 10,000–8,000 BCE.

The Southern African hunter-gatherers (Khoisan) are suggested to represent the autochthonous hunter-gatherer population of southern Africa, prior to the expansion of Bantu-speakers from Western/Central Africa and East African pastoralists. Khoisan show evidence for Bantu-related admixture, ranging from nearly ~0% to up to ~87.1%.

Out-of-Africa event

Human migration routes following Out-of-Africa.
Most modern Africans display a high level of genetic homogeneity, but contributions from Eurasian populations are substantial, mostly concentrated in the Northeastern part of Africa and Madagascar.

The "recent African origin of modern humans" proposes a "single origin" of Homo sapiens within Africa. Recent genetic and archeologic data suggests that Homo sapiens-subgroups originated in multiple regions of Africa, not confined to a single sub-region of origin, with the last common ancestor of all modern humans expanding from a single region absorbing or replacing various deep lineages (described as archaic ghosts). The H. sapiens ancestral to proper Eurasians most likely left Northeastern Africa between 50,000 and 100,000 years ago. The "recent African origin" model proposes that all modern non-African populations descend from one or several waves of H. sapiens that left Africa 70,000–60,000 years ago.

According to serial founder model, the earliest-branching non-African lineages are expected to have expanded in the Middle East, after the Out-of-Africa event (a), but have their deepest divergence in modern-day East or Southeast Asia (b), suggesting either rapid diversification and substructure within the early Eurasians, or replacement and loss of deep lineages in Western Eurasia. Simplified Y tree is shown as reference for colours.

According to Durvasula et al. (2020), there are indications that 2% to 19% (≃6.6 to 7.0%) of the DNA of West African populations may have come from an unknown archaic hominin which split from the ancestor of humans and Neanderthals between 360 kya to 1.02 mya. However, Durvasula et al. (2020) also suggests that at least part of this archaic admixture is also present in Eurasians/non-Africans, and that the admixture event or events range from 0 to 124 ka B.P, which includes the period before the Out-of-Africa migration and prior to the African/Eurasian split (thus affecting in part the common ancestors of both Africans and Eurasians/non-Africans). Chen et al. (2020) found that Africans have higher Neanderthal ancestry than previously thought. 2,504 African samples from all over Africa were analyzed and tested on Neanderthal ancestry. All African samples showed evidence for minor Neanderthal ancestry, but always at lower levels than observed in Eurasians.

Geneflow between Eurasian and African populations

Pre-Neolithic and Neolithic migration events in Africa.
A visual summary of the main admixture movements into and within Africa.

Significant Eurasian admixture is found in Northern Africa, and among specific ethnic groups of the Horn of Africa, Northern Sudan, the Sahel region, as well as among the Malagasy people of Madagascar. Various genome studies found evidence for multiple prehistoric back-migrations from various Eurasian populations and subsequent admixture with native groups. West Eurasian-associated geneflow arrived to Northern Africa during the Paleolithic (30,000 to 15,000 years ago), followed by other pre-Neolithic and Neolithic migration events. Genetic data on the Taforalt samples "demonstrated that Northern Africa received significant amounts of gene-flow from Eurasia predating the Holocene and development of farming practices". Medieval geneflow events, such as the Arab expansion also left traces in various African populations. Pickrell et al. (2014) indicated that Western Eurasian ancestry eventually arrived through Northeast Africa (particularly the Horn of Africa) to Southeast Africa and Southern Africa.

Ramsay et al. (2018) also found evidence for significant Western Eurasian admixture in various parts of Africa, from both ancient and more recent migrations, being highest among populations from Northern Africa, and some groups of the Horn of Africa:

In addition to the intrinsic diversity within the continent due to population structure and isolation, migration of Eurasian populations into Africa has emerged as a critical contributor to the genetic diversity. These migrations involved the influx of different Eurasian populations at different times and to different parts of Africa. Comprehensive characterization of the details of these migrations through genetic studies on existing populations could help to explain the strong genetic differences between some geographically neighbouring populations.

This distinctive Eurasian admixture appears to have occurred over at least three time periods with ancient admixture in central west Africa (e.g., Yoruba from Nigeria) occurring between ~7.5 and 10.5 kya, older admixture in east Africa (e.g., Ethiopia) occurring between ~2.4 and 3.2 kya and more recent admixture between ~0.15 and 1.5 kya in some east African (e.g., Kenyan) populations.

Subsequent studies based on LD decay and haplotype sharing in an extensive set of African and Eurasian populations confirmed the presence of Eurasian signatures in west, east and southern Africans. In the west, in addition to Niger-Congo speakers from The Gambia and Mali, the Mossi from Burkina Faso showed the oldest Eurasian admixture event ~7 kya. In the east, these analyses inferred Eurasian admixture within the last 4000 years in Kenya.

Ancestral components of various human populations using a four-way Admixture model. European and Middle Eastern (Western Eurasian) ancestry is found in many African groups.

There is no definitive agreement on when or where the original homeland of the Afroasiatic language family existed. Some have suggested that they were spread by people with largely West-Eurasian ancestry during the Neolithic Revolution, towards Northern Africa and the Horn of Africa, outgoing from the Middle East, specifically from the Levant. Others argue that the first speakers of Proto-Afroasiatic were based in Northeast Africa because that region includes the majority of the diversity of the Afroasiatic language family and has very diverse groups in close geographic proximity, which is sometimes considered a telltale sign for a linguistic geographic origin. A subset of the Proto-Afroasiatic population would have migrated to the Levant during the late Paleolithic, merging with local West-Eurasians and resulting in a population which would later give rise to Natufian culture, associated with the early development of agriculture and early Afroasiatic languages, or specifically pre-proto-Semitic. In addition, Y-haplogroup sub-lineage E-M215 (also known as "E1b1b) and its derivative E-M35 are quite common among Afroasiatic speakers, and southwestern Ethiopia is a plausible source of these haplogroups. Under this African model, the linguistic group and carriers of this lineage would have arisen and dispersed together from Northeast Africa in the Mesolithic, plausibly having already developed subsistence patterns of pastoralism and intensive plant usage and collection.

The Near-Eastern agriculturalist hypothesis does not account for the domestication of plants endemic to the Horn of Africa such as teff, ensete, and Niger seed, nor does it account for the lack of evidence of intrusive agricultural populations or for the growing of wheat, barley, or sorghum in that region prior to 3000 B.C. According to historian and linguist Christopher Ehret, the form of intensive plant collection practiced by the Proto-Afroasiatic population in Northeast Africa may have been a precursor to the other agricultural practices that would later independently develop in the Fertile Crescent and the Horn of Africa.

Proposed migration routes of paternal lineage E.

Horn of Africa

While many studies conducted on Horn of Africa populations estimate a West-Eurasian admixture event around 3,000 years ago, Hodgson et al. (2014) found a distinct West-Eurasian ancestral component among studied Afroasiatic-speaking groups in the Horn of Africa (and to a lesser extent in North Africa and West Asia), most prevalent among the ethnic Somali. This ancestral component dubbed "Ethio-Somali" is most closely related to the "Maghrebi" (peaking in Tunisians) component and is believed to have diverged from other non-African ancestries around 23,000 years ago, and migrated back to Africa prior to developing agriculture (12–23 ka) from the Near East. This population would have crossed via the Sinai Peninsula and then split into two, with one branch continuing west across North Africa and the other heading south into the Horn of Africa. The authors propose that the "Ethio-Somali" component may have been a substantial ancestral component of the Proto-Afroasiatic-speaking population. Later migration from Arabia into the HOA beginning around 3 ka would explain the origin of the Ethiosemitic languages at this time. An mtDNA analysis by Gandini et al. (2016) has produced additional evidence in support of a pre-agricultural back-migration from West-Eurasia into the Horn of Africa with an estimated date of arrival into the Horn of Africa in the early Holocene, possibly as a result of obsidian exchange networks across the Red Sea. Hodgson et al. also confirmed the existence of an ancestral component indigenous to the Horn of Africa - "Ethiopic" or "Omotic" (Pagani et al.) - which is most prevalent among speakers of the Omotic branch of Afroasiatic in southwestern Ethiopia. This lineage is associated with that of a 4,500 year-old fossil (Mota) found in a cave in southwestern Ethiopia, which has high genetic affinity to modern Ethiopian groups, especially the endogamous blacksmith caste of the Omotic Aari people. Like Mota, Aari blacksmiths do not show evidence for admixture with West-Eurasians, demonstrating a degree of population continuity in this region for at least 4,500 years. In a comparative analysis of Mota's genome referencing modern populations, Gallego et al. (2016) concluded that the divergence of Omotic from other Afroasiatic languages may have resulted from the relative isolation of its speakers from external groups.

In an analysis of 68 Ethiopian ethnic groups, Lopez et al. (2021) revealed that several groups belonging to the three AA classifications of Cushitic, Omotic and Semitic show high genetic similarity to each other on average. Furthermore, the Nilo-Saharan speakers in the southwest shared more recent ancestry with Bantu and Nilotics, in contrast Afro-Asiatic speakers in the northeast shared more recent ancestry with Egyptians and other West Eurasians. The data also supported widespread recent intermixing among various ethnic groups.

Madagascar

Austronesian expansion, outgoing from Taiwan and the northern Philippines.

Specific East Asian-related ancestry is found among the Malagasy speakers of Madagascar at a medium frequency. The presence of this East Asian-related ancestry is mostly linked to the Austronesian peoples expansion from Southeast Asia. The peoples of Borneo were identified to resemble the East Asian voyagers, who arrived on Madagascar. East Asian ancestry among Malagasy people was estimated at a mean average of 33%, but as high as ~75% among some Highlander groups and upper caste groups.

Northern Africa

Dobon et al. (2015) identified an autosomal ancestral component that is commonly found among modern Afroasiatic-speaking populations (as well as Nubians) in Northeast Africa. This Coptic component peaks among Copts in Sudan, which is differentiated by its lack of Arab influence, but shares common ancestry with the North African/Middle Eastern populations. It appears alongside a component that defines Nilo-Saharan speakers of southwestern Sudan and South Sudan. Arauna et al. (2017), analyzing existing genetic data obtained from Northern African populations, such as Berbers, described them as a mosaic of North African (Taforalt), Middle Eastern, European (Early European Farmers), and Sub-Saharan African-related ancestries.

Chen et al. (2020) analyzed 2,504 African samples from all over Africa, and found archaic Neanderthal ancestry, among all tested African samples at low frequency. They also identified a European-related (West-Eurasian) ancestry segment, which seems to largely correspond with the detected Neanderthal ancestry components. European-related admixture among Africans was estimated to be between ~0% to up to ~30%, with a peak among Northern Africans. According to Chen et al. (2020), "These data are consistent with the hypothesis that back-migration contributed to the signal of Neanderthal ancestry in Africans. Furthermore, the data indicates that this back-migration came after the split of Europeans and East Asians, from a population related to the European lineage."

There is a minor geneflow from North Africa in parts of Southern Europe, this is supported by the presence of an African-specific mitochondrial haplogroup among one of four 4,000 year old samples. Multiple studies found also evidence for geneflow of African ancestry towards Eurasia, specifically Europe and the Middle East. The analysis of 40 different West-Eurasian populations found African admixture at a frequency of 0% to up to ~15%.

Western Africa

Hollfelder et al. (2021) concluded that West African Yoruba people, which were previously used as "unadmixed reference population" for indigenous Africans, harbor minor levels of Neanderthal ancestry, which can be largely associated with back-migration of an "Ancestral European-like" source population.

A genome-wide study of a Fulani community from Burkina Faso inferred two major admixture events in this group, dating to ~1800 ya, and 300 ya. The first admixture event took place between the West African ancestors of the Fula and ancestral North African nomadic groups. The second admixture event, relatively recent, inferred a source from Southwestern Europe, or suggests either an additional gene flow between the Fulani and Northern African groups, who carry admixture proportions from Europeans. Sahelian populations like the Toubou also showed admixture coming from Eurasians.

Southern Africa

Low levels of West Eurasian ancestry (European or Middle Eastern) are found in Khoe–Kwadi Khoesan-speakers. It could have been acquired indirectly by admixture with migrating pastoralists from East Africa. This hypothesis of gene flow from eastern to southern Africa is further supported by other genetic and archaeological data documenting the spread of pastoralism from East to South Africa.

Regional genomic overview

North Africa

Archaic Human DNA

While Denisovan and Neanderthal ancestry in non-Africans outside of Africa are more certain, archaic human ancestry in Africans is less certain and is too early to be established with certainty.

Ancient DNA

Daniel Shriner (2018), using modern populations as a reference, showed that the Natufians carried 61.2% Arabian, 21.2% Northern African, 10.9% Western Asian, and a small portion of Eastern African ancestry at 6.8%, which is associated with the modern Omotic-speaking groups found in southern Ethiopia.

Egypt

Khnum-aa, Khnum-Nakht, Nakht-Ankh and JK2911 carried maternal haplogroup M1a1.

Djehutynakht (10A) carried maternal haplogroup U5b2b5. JK2888 carried maternal haplogroup U6a2.

Thuya, Tiye, Tutankhamen's mother, and Tutankhamen carried the maternal haplogroup K.

JK2134 carried maternal haplogroup J1d and JK2887 carried maternal haplogroup J2a1a1.

Amenhotep III, Akhenaten, and Tutankhamen carried the paternal haplogroup R1b.

Ramesses III and "Unknown Man E", possibly Pentawere, carried paternal haplogroup E1b1a.

JK2134 and JK2911 carried paternal haplogroup J.

Takabuti carried maternal haplogroup H4a1 and YM:KMM A 63 carried maternal haplogroup HV.

OM:KMM A 64 carried maternal haplogroup T2c1a.

JK2888 carried paternal haplogroup E1b1b1a1b2.

Libya

At Takarkori rockshelter, in Libya, two naturally mummified women, dated to the Middle Pastoral Period (7000 BP), carried basal maternal haplogroup N.

Morocco

Van de Loorsdrecht et al. (2018) found that of seven samples of Taforalts of Morocco, radiocarbon dated to between 15,100 cal BP and 13,900 cal BP, six were found to carry maternal haplogroup U6a, and one was found to carry maternal haplogroup M1b. All six males were found to carry paternal haplogroup E1b1b, and they harbored 63.5% Natufian-related ancestry and 36.5% Sub-Saharan African-related ancestry. The Sub-Saharan component is most strongly drawn out by modern West African groups such as the Yoruba and the Mende. The samples also contain an additional affinity to South, Central, and East African outgroups that cannot be explained by any known ancient or modern populations. When projected onto a principal component analysis graph of African and west Eurasian populations, the Taforalt individuals form a distinct cluster in an intermediate position between modern North Africans (e.g., Berbers, Mozabites, Saharawis) and East Africans (e.g., Afars, Oromos, Somalis). Jeong (2020), when comparing the Taforalt people of the Iberomaurusian culture to modern populations, found that the Taforalt's Sub-Saharan African genetic component may be best represented by modern West Africans (e.g., Yoruba).

Y-Chromosomal DNA

Mitochondrial DNA

Amid the Holocene, including the Holocene Climate Optimum in 8000 BP, Africans bearing haplogroup L2 spread within West Africa and Africans bearing haplogroup L3 spread within East Africa. As the largest migration since the Out of Africa migration, migration from Sub-Saharan Africa toward the North Africa occurred, by West Africans, Central Africans, and East Africans, resulting in migrations into Europe and Asia; consequently, Sub-Saharan African mitochondrial DNA was introduced into Europe and Asia. During the early period of the Holocene, 50% of Sub-Saharan African mitochondrial DNA was introduced into North Africa by West Africans and the other 50% was introduced by East Africans. During the modern period, a greater number of West Africans introduced Sub-Saharan African mitochondrial DNA into North Africa than East Africans.

Mitochondrial haplogroups L3, M, and N are found among Sudanese peoples (e.g., Beja, Nilotics, Nuba, Nubians), who have no known interaction (e.g., history of migration/admixture) with Europeans or Asians; rather than having developed in a post-Out-of-Africa migration context, mitochondrial macrohaplogroup L3/M/N and its subsequent development into distinct mitochondrial haplogroups (e.g., Haplogroup L3, Haplogroup M, Haplogroup N) may have occurred in East Africa at a time that considerably predates the Out-of-Africa migration event of 50,000 BP.

Autosomal DNA

Medical DNA

The genomes of Africans commonly found to undergo adaptation are regulatory DNA, and many cases of adaptation found among Africans relate to diet, physiology, and evolutionary pressures from pathogens.

Lactase persistence

Neolithic agriculturalists, who may have resided in Northeast Africa and the Near East, may have been the source population for lactase persistence variants, including –13910*T, and may have been subsequently supplanted by later migrations of peoples. The Sub-Saharan West African Fulani, the North African Tuareg, and European agriculturalists, who are descendants of these Neolithic agriculturalists, share the lactase persistence variant –13910*T. While shared by Fulani and Tuareg herders, compared to the Tuareg variant, the Fulani variant of –13910*T has undergone a longer period of haplotype differentiation. The Fulani lactase persistence variant –13910*T may have spread, along with cattle pastoralism, between 9686 BP and 7534 BP, possibly around 8500 BP; corroborating this timeframe for the Fulani, by at least 7500 BP, there is evidence of herders engaging in the act of milking in the Central Sahara.

West Africa

Archaic Human DNA

Archaic traits found in human fossils of West Africa (e.g., Iho Eleru fossils, which dates to 13,000 BP) and Central Africa (e.g., Ishango fossils, which dates between 25,000 BP and 20,000 BP) may have developed as a result of admixture between archaic humans and modern humans or may be evidence of late-persisting early modern humans. While Denisovan and Neanderthal ancestry in non-Africans outside of Africa are more certain, archaic human ancestry in Africans is less certain and is too early to be established with certainty.

Ancient DNA

As of 2017, human ancient DNA has not been found in the region of West Africa. As of 2020, human ancient DNA has not been forthcoming in the region of West Africa.

Y-Chromosomal DNA

Eight male individuals from Guinea Bissau, two male individuals from Niger, one male individual from Mali, and one male individual from Cabo Verde carried haplogroup A1a.

As a result of haplogroup D0, a basal branch of haplogroup DE, being found in three Nigerian men, it may be the case that haplogroup DE, as well as its sublineages D0 and E, originated in Africa.

As of 19,000 years ago, Africans, bearing haplogroup E1b1a-V38, likely traversed across the Sahara, from east to westE1b1a1-M2 likely originated in West Africa or Central Africa.

Mitochondrial DNA

Around 18,000 BP, Mende people, along with Gambian peoples, grew in population size.

In 15,000 BP, Niger-Congo speakers may have migrated from the Sahelian region of West Africa, along the Senegal River, and introduced L2a1 into North Africa, resulting in modern Mauritanian peoples and Berbers of Tunisia inheriting it.

Between 11,000 BP and 10,000 BP, Yoruba people and Esan people grew in population size.

As early as 11,000 years ago, Sub-Saharan West Africans, bearing macrohaplogroup L (e.g., L1b1a11, L1b1a6a, L1b1a8, L1b1a9a1, L2a1k, L3d1b1a), may have migrated through North Africa and into Europe, mostly into southern Europe (e.g., Iberia).

Autosomal DNA

During the early period of the Holocene, in 9000 BP, Khoisan-related peoples admixed with the ancestors of the Igbo people, possibly in the western Sahara.

Between 2000 BP and 1500 BP, Nilo-Saharan-speakers may have migrated across the Sahel, from East Africa into West Africa, and admixed with Niger-Congo-speaking Berom people. In 710 CE, West African-related populations (e.g., Niger-Congo-speaking Berom people, Bantu-speakers) and East African-related populations (Nilo-Saharan-speaking Ethiopians, Nilo-Saharan-speaking Chadians) admixed with one another in northern Nigeria and northern Cameroon.

Fan et al. (2019) found that the Fulani people show genetic affinity to isolated Afroasiatic-speaking groups in Eastern Africa, specifically Omotic-speakers such as the Aari people. While the Fulani have nearly exclusive indigenous African ancestry (defined by West and East African ancestry), they also show traces of West-Eurasian-like admixture, supporting an ancestral homeland somewhere in North or Eastern Africa, and westwards expansion during the Neolithic, possibly caused by the arrival and expansion of West-Eurasian-related groups. Fan et al. (2023) found that the Fulani, who have 50% Amhara-related and 50% Tikari-related ancestry as well as occupy regions such as West Africa, Central Africa, and the Sudan as nomadic herders, may have initially been Afroasiatic speakers that subsequently underwent language replacement and became Niger-Congo speakers.

Medical DNA

The genomes of Africans commonly found to undergo adaptation are regulatory DNA, and many cases of adaptation found among Africans relate to diet, physiology, and evolutionary pressures from pathogens.[92] Throughout Sub-Saharan Africa, genetic adaptation (e.g., rs334 mutation, Duffy blood group, increased rates of G6PD deficiency, sickle cell disease) to malaria has been found among Sub-Saharan Africans, which may have initially developed in 7300 BP.[92] Sub-Saharan Africans have more than 90% of the Duffy-null genotype.[108]

Pediculus

During the Copper Age and early Islamic era of ancient Israel, West Africans may have migrated into ancient Israel and introduced head louse from West Africa.

Sickle Cell

Amid the Green Sahara, the mutation for sickle cell originated in the Sahara or in the northwest forest region of western Central Africa (e.g., Cameroon) by at least 7,300 years ago, though possibly as early as 22,000 years ago. The ancestral sickle cell haplotype to modern haplotypes (e.g., Cameroon/Central African Republic and Benin/Senegal haplotypes) may have first arose in the ancestors of modern West Africans, bearing haplogroups E1b1a1-L485 and E1b1a1-U175 or their ancestral haplogroup E1b1a1-M4732. West Africans (e.g., Yoruba and Esan of Nigeria), bearing the Benin sickle cell haplotype, may have migrated through the northeastern region of Africa into the western region of Arabia. West Africans (e.g., Mende of Sierra Leone), bearing the Senegal sickle cell haplotype, may have migrated into Mauritania (77% modern rate of occurrence) and Senegal (100%); they may also have migrated across the Sahara, into North Africa, and from North Africa, into Southern Europe, Turkey, and a region near northern Iraq and southern Turkey. Some may have migrated into and introduced the Senegal and Benin sickle cell haplotypes into Basra, Iraq, where both occur equally. West Africans, bearing the Benin sickle cell haplotype, may have migrated into the northern region of Iraq (69.5%), Jordan (80%), Lebanon (73%), Oman (52.1%), and Egypt (80.8%).

Schistosomes

According to Steverding (2020), while not definite: Near the African Great Lakes, schistosomes (e.g., S. mansoni, S. haematobium) underwent evolution. Subsequently, there was an expansion alongside the Nile. From Egypt, the presence of schistosomes may have expanded, via migratory Yoruba people, into Western Africa. Thereafter, schistosomes may have expanded, via migratory Bantu peoples, into the rest of Sub-Saharan Africa (e.g., Southern Africa, Central Africa).

Thalassemia

Through pathways taken by caravans, or via travel amid the Almovarid period, a population (e.g., Sub-Saharan West Africans) may have introduced the –29 (A → G) β-thalassemia mutation (found in notable amounts among African Americans) into the North African region of Morocco.

Domesticated Animal DNA

While the Niger-Congo migration may have been from West Africa into Kordofan, possibly from Kordofan, Sudan, Niger-Congo speakers accompanied by undomesticated helmeted guineafowls, may have traversed into West Africa, domesticated the helmeted guineafowls by 3000 BCE, and via the Bantu expansion, traversed into other parts of Sub-Saharan Africa (e.g., Central Africa, East Africa, Southern Africa).

Central Africa

Archaic Human DNA

Archaic traits found in human fossils of West Africa (e.g., Iho Eleru fossils, which dates to 13,000 BP) and Central Africa (e.g., Ishango fossils, which dates between 25,000 BP and 20,000 BP) may have developed as a result of admixture between archaic humans and modern humans or may be evidence of late-persisting early modern humans. While Denisovan and Neanderthal ancestry in non-Africans outside of Africa are more certain, archaic human ancestry in Africans is less certain and is too early to be established with certainty.

Ancient DNA

In 4000 BP, there may have been a population that traversed from Africa (e.g., West Africa or West-Central Africa), through the Strait of Gibraltar, into the Iberian Peninsula, where admixing between Africans and Iberians (e.g., of northern Portugal, of southern Spain) occurred.

Cameroon

West African hunter-gatherers, in the region of western Central Africa (e.g., Shum Laka, Cameroon), particularly between 8000 BP and 3000 BP, were found to be related to modern Central African hunter-gatherers (e.g., Baka, Bakola, Biaka, Bedzan).

Democratic Republic of Congo

At Kindoki, in the Democratic Republic of Congo, there were three individuals, dated to the protohistoric period (230 BP, 150 BP, 230 BP); one carried haplogroups E1b1a1a1d1a2 (E-CTS99, E-CTS99) and L1c3a1b, another carried haplogroup E (E-M96, E-PF1620), and the last carried haplogroups R1b1 (R-P25 1, R-M415) and L0a1b1a1.

Y-Chromosomal DNA

Haplogroup R1b-V88 is thought to have originated in Europe and migrated into Africa with farmers or herders in the Neolithic period, c. 5500 BC. R1b-V88 is found at a high frequency among Chadic speaking peoples such as the Hausa, as well as in KanembuFulani, and Toubou populations.

Mitochondrial DNA

In 150,000 BP, Africans (e.g., Central Africans, East Africans) bearing haplogroup L1 diverged. Between 75,000 BP and 60,000 BP, Africans bearing haplogroup L3 emerged in East Africa and eventually migrated into and became present in modern West Africans, Central Africans, and non-Africans. Amid the Holocene, including the Holocene Climate Optimum in 8000 BP, Africans bearing haplogroup L2 spread within West Africa and Africans bearing haplogroup L3 spread within East Africa. As the largest migration since the Out of Africa migration, migration from Sub-Saharan Africa toward the North Africa occurred, by West Africans, Central Africans, and East Africans, resulting in migrations into Europe and Asia; consequently, Sub-Saharan African mitochondrial DNA was introduced into Europe and Asia.

Mitochondrial haplogroup L1c is strongly associated with pygmies, especially with Bambenga groups. L1c prevalence was variously reported as: 100% in Ba-Kola, 97% in Aka (Ba-Benzélé), and 77% in Biaka, 100% of the Bedzan (Tikar), 97% and 100% in the Baka people of Gabon and Cameroon, respectively, 97% in Bakoya (97%), and 82% in Ba-Bongo. Mitochondrial haplogroups L2a and L0a are prevalent among the Bambuti.

Autosomal DNA

Genetically, African pygmies have some key difference between them and Bantu peoples.

Medical DNA

Evidence suggests that, when compared to other Sub-Saharan African populations, African pygmy populations display unusually low levels of expression of the genes encoding for human growth hormone and its receptor associated with low serum levels of insulin-like growth factor-1 and short stature.

The genomes of Africans commonly found to undergo adaptation are regulatory DNA, and many cases of adaptation found among Africans relate to diet, physiology, and evolutionary pressures from pathogens. Throughout Sub-Saharan Africa, genetic adaptation (e.g., rs334 mutation, Duffy blood group, increased rates of G6PD deficiency, sickle cell disease) to malaria has been found among Sub-Saharan Africans, which may have initially developed in 7300 BP. Sub-Saharan Africans have more than 90% of the Duffy-null genotype. In the rainforests of Central Africa, genetic adaptation for non-height-related factors (e.g., immune traits, reproduction, thyroid function) and short stature (e.g., EHB1 and PRDM5 – bone synthesis; OBSCN and COX10 – muscular development; HESX1 and ASB14 – pituitary gland's growth hormone production/secretion) has been found among rainforest hunter-gatherers.

Eastern Africa

From the region of Kenya and Tanzania to South Africa, eastern Bantu-speaking Africans constitute a north to south genetic cline; additionally, from eastern Africa to toward southern Africa, evidence of genetic homogeneity is indicative of a serial founder effect and admixture events having occurred between Bantu-speaking Africans and other African populations by the time the Bantu migration had spanned into South Africa.

Archaic Human DNA

While Denisovan and Neanderthal ancestry in non-Africans outside of Africa are more certain, archaic human ancestry in Africans is less certain and is too early to be established with certainty.

Ancient DNA

Ethiopia

At Mota, in Ethiopia, an individual, estimated to date to the 5th millennium BP, carried haplogroups E1b1 and L3x2a. The individual of Mota is genetically related to groups residing near the region of Mota, and in particular, are considerably genetically related to the Aari people, especially the blacksmith caste of that group.

Kenya

At Jawuoyo Rockshelter, in Kisumu County, Kenya, a forager of the Later Stone Age carried haplogroups E1b1b1a1b2/E-V22 and L4b2a2c.

At Ol Kalou, in Nyandarua County, Kenya, a pastoralist of the Pastoral Neolithic carried haplogroups E1b1b1b2b2a1/E-M293 and L3d1d.

At Kokurmatakore, in Marsabit County, Kenya, a pastoralist of the Pastoral Iron Age carried haplogroups E1b1b1/E-M35 and L3a2a.

At White Rock Point, in Homa Bay County, Kenya, there were two foragers of the Later Stone Age; one carried haplogroups BT (xCT), likely B, and L2a4, and another probably carried haplogroup L0a2.

At Nyarindi Rockshelter, in Kenya, there were two individuals, dated to the Later Stone Age (3500 BP); one carried haplogroup L4b2a and another carried haplogroup E (E-M96, E-P162).

At Lukenya Hill, in Kenya, there were two individuals, dated to the Pastoral Neolithic (3500 BP); one carried haplogroups E1b1b1b2b (E-M293, E-CTS10880) and L4b2a2b, and another carried haplogroup L0f1.

At Hyrax Hill, in Kenya, an individual, dated to the Pastoral Neolithic (2300 BP), carried haplogroups E1b1b1b2b (E-M293, E-M293) and L5a1b.

At Molo Cave, in Kenya, there were two individuals, dated to the Pastoral Neolithic (1500 BP); while one had haplogroups that went undetermined, another carried haplogroups E1b1b1b2b (E-M293, E-M293) and L3h1a2a1.

At Kakapel, in Kenya, there were three individuals, one dated to the Later Stone Age (3900 BP) and two dated to the Later Iron Age (300 BP, 900 BP); one carried haplogroups CT (CT-M168, CT-M5695) and L3i1, another carried haplogroup L2a1f, and the last carried haplogroup L2a5.

At Panga ya Saidi, in Kenya, an individual, estimated to date between 496 BP and 322 BP, carried haplogroups E1b1b1b2 and L4b2a2.

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1250 CE and 1650 CE, carried haplogroup L3b1a1a.

Tanzania

At Mlambalasi rockshelter, in Tanzania, an individual, dated between 20,345 BP and 17,025 BP, carried undetermined haplogroups.

At Gishimangeda Cave, in Karatu District, Tanzania, there were eleven pastoralists of the Pastoral Neolithic; one carried haplogroups E1b1b1a1b2/E-V22 and HV1b1, another carried haplogroup L0a, another carried haplogroup L3x1, another carried haplogroup L4b2a2b, another carried haplogroups E1b1b1b2b2a1/E-M293 and L3i2, another carried haplogroup L3h1a2a1, another carried haplogroups E1b1b1b2b2/E-V1486, likely E-M293 and L0f2a1, and another carried haplogroups E1b1b1b2b2/E-V1486, likely E-M293, and T2+150; while most of the haplogroups among three pastoralists went undetermined, one was determined to carry haplogroup BT, likely B.

At Kilwa, Coast, in Tanzania, an individual, dated between 1300 CE and 1600 CE, carried haplogroups J2a2a1a1a2a~ and L2a1h.

At Lindi, in Tanzania, an individual, dated between 1511 cal CE and 1664 cal CE, carried haplogroups E1b1a1a1a2a1a3a1d~ and L0a1a2.

At Makangale Cave, on Pemba Island, Tanzania, an individual, estimated to date between 1421 BP and 1307 BP, carried haplogroup L0a.

At Songo Mnara, in Tanzania, an individual, dated between 1294 cal CE and 1392 cal CE, carried haplogroups R1a and L3e3a.

Uganda

At Munsa, in Uganda, an individual, dated to the Later Iron Age (500 BP), carried haplogroup L3b1a1.

Y-Chromosomal DNA

As of 19,000 years ago, Africans, bearing haplogroup E1b1a-V38, likely traversed across the Sahara, from east to west.

Before the slave trade period, East Africans, who carried haplogroup E1b1a-M2, expanded into Arabia, resulting in various rates of inheritance throughout Arabia (e.g., 2.8% Qatar, 3.2% Yemen, 5.5% United Arab Emirates, 7.4% Oman).

Mitochondrial DNA

In 150,000 BP, Africans (e.g., Central Africans, East Africans) bearing haplogroup L1 diverged. In 130,000 BP, Africans bearing haplogroup L5 diverged in East Africa. Between 130,000 BP and 75,000 BP, behavioral modernity emerged among Southern Africans and long-term interactions between the regions of Southern Africa and Eastern Africa became established. Between 75,000 BP and 60,000 BP, Africans bearing haplogroup L3 emerged in East Africa and eventually migrated into and became present in modern West Africans, Central Africans, and non-Africans. Amid the Holocene, including the Holocene Climate Optimum in 8000 BP, Africans bearing haplogroup L2 spread within West Africa and Africans bearing haplogroup L3 spread within East Africa. As the largest migration since the Out of Africa migration, migration from Sub-Saharan Africa toward the North Africa occurred, by West Africans, Central Africans, and East Africans, resulting in migrations into Europe and Asia; consequently, Sub-Saharan African mitochondrial DNA was introduced into Europe and Asia. During the early period of the Holocene, 50% of Sub-Saharan African mitochondrial DNA was introduced into North Africa by West Africans and the other 50% was introduced by East Africans. During the modern period, a greater number of West Africans introduced Sub-Saharan African mitochondrial DNA into North Africa than East Africans. Between 15,000 BP and 7000 BP, 86% of Sub-Saharan African mitochondrial DNA was introduced into Southwest Asia by East Africans, largely in the region of Arabia, which constitute 50% of Sub-Saharan African mitochondrial DNA in modern Southwest Asia. In the modern period, 68% of Sub-Saharan African mitochondrial DNA was introduced by East Africans and 22% was introduced by West Africans, which constitutes 50% of Sub-Saharan African mitochondrial DNA in modern Southwest Asia.

Autosomal DNA

Across all areas of Madagascar, the average ancestry for the Malagasy people was found to be 4% West Eurasian, 37% Austronesian, and 59% Bantu.

Medical DNA

The genomes of Africans commonly found to undergo adaptation are regulatory DNA, and many cases of adaptation found among Africans relate to diet, physiology, and evolutionary pressures from pathogens. Throughout Sub-Saharan Africa, genetic adaptation (e.g., rs334 mutation, Duffy blood group, increased rates of G6PD deficiency, sickle cell disease) to malaria has been found among Sub-Saharan Africans, which may have initially developed in 7300 BP. Sub-Saharan Africans have more than 90% of the Duffy-null genotype. In the highlands of Ethiopia, genetic adaptation (e.g., rs10803083, an SNP associated with the rate and function of hemoglobin; BHLHE41, a gene associated with circadian rhythm and hypoxia response; EGNL1, a gene strongly associated with oxygen homeostasis in mammals) to hypoxia and low atmospheric pressure has been found among the Amhara people, which may have developed within the past 5000 years. In Tanzania, genetic adaptation (e.g., greater amount of amylase genes than in African populations that consume low-starch foods) has been found in the Hadza people due to a food diet that especially includes consumption of tubers.

Southern Africa

From the region of Kenya and Tanzania to South Africa, eastern Bantu-speaking Africans constitute a north to south genetic cline; additionally, from eastern Africa to toward southern Africa, evidence of genetic homogeneity is indicative of a serial founder effect and admixture events having occurred between Bantu-speaking Africans and other African populations by the time the Bantu migration had spanned into South Africa.

Archaic Human DNA

While Denisovan and Neanderthal ancestry in non-Africans outside of Africa are more certain, archaic human ancestry in Africans is less certain and is too early to be established with certainty.

Ancient DNA

Three Later Stone Age hunter-gatherers carried ancient DNA similar to Khoisan-speaking hunter-gatherers. Prior to the Bantu migration into the region, as evidenced by ancient DNA from Botswana, East African herders migrated into Southern Africa. Out of four Iron Age Bantu agriculturalists of West African origin, two earlier agriculturalists carried ancient DNA similar to Tsonga and Venda peoples and the two later agriculturalists carried ancient DNA similar to Nguni people; this indicates that there were various movements of peoples in the overall Bantu migration, which resulted in increased interaction and admixing between Bantu-speaking peoples and Khoisan-speaking peoples.

Botswana

At Nqoma, in Botswana, an individual, dated to the Early Iron Age (900 BP), carried haplogroup L2a1f.

At Taukome, in Botswana, an individual, dated to the Early Iron Age (1100 BP), carried haplogroups E1b1a1 (E-M2, E-Z1123) and L0d3b1.

At Xaro, in Botswana, there were two individuals, dated to the Early Iron Age (1400 BP); one carried haplogroups E1b1a1a1c1a and L3e1a2, and another carried haplogroups E1b1b1b2b (E-M293, E-CTS10880) and L0k1a2.

Malawi

At Fingira rockshelter, in Malawi, an individual, dated between 6179 BP and 2341 BP, carried haplogroups B2 and L0d1.

At Chencherere, in Malawi, an individual, estimated to date between 5400 BP and 4800 BP, carried haplogroup L0k2.

At Hora 1 rockshelter, in Malawi, an individual, dated between 16,897 BP and 15,827 BP, carried haplogroups B2b and L5b.

South Africa

At Doonside, in South Africa, an individual, estimated to date between 2296 BP and 1910 BP, carried haplogroup L0d2.

At Ballito Bay, South Africa, an individual, estimated to date between 1986 BP and 1831 BP, carried haplogroups A1b1b2 and L0d2c1.

At Kalemba rockshelter, in Zambia, an individual, dated between 5285 BP and 4975 BP, carried haplogroup L0d1b2b.

Y-Chromosomal DNA

Distribution of Y-Chromosome Haplogroup A in Africa.
Distribution of haplogroup B (M60) of the human Y chromosome in native populations.

Various Y chromosome studies show that the San carry some of the most divergent (oldest) human Y-chromosome haplogroups. These haplogroups are specific sub-groups of haplogroups A and B, the two earliest branches on the human Y-chromosome tree.

Mitochondrial DNA

In 200,000 BP, Africans (e.g., Khoisan of Southern Africa) bearing haplogroup L0 diverged from other Africans bearing haplogroup L1′6, which tend to be northward of Southern Africa. Between 130,000 BP and 75,000 BP, behavioral modernity emerged among Southern Africans and long-term interactions between the regions of Southern Africa and Eastern Africa became established.

Mitochondrial DNA studies also provide evidence that the San carry high frequencies of the earliest haplogroup branches in the human mitochondrial DNA tree. This DNA is inherited only from one's mother. The most divergent (oldest) mitochondrial haplogroup, L0d, has been identified at its highest frequencies in the southern African San groups.

Autosomal DNA

Henn et al. (2011) found that the ǂKhomani San, as well as the Sandawe and Hadza peoples of Tanzania, were the most genetically diverse of any living humans studied. This high degree of genetic diversity hints at the origin of anatomically modern humans.

Medical DNA

Among the ancient DNA from three hunter-gatherers sharing genetic similarity with San people and four Iron Age agriculturalists, their SNPs indicated that they bore variants for resistance against sleeping sickness and Plasmodium vivax. In particular, two out of the four Iron Age agriculturalists bore variants for resistance against sleeping sickness and three out of the four Iron Age agriculturalists bore Duffy negative variants for resistance against malaria. In contrast to the Iron Age agriculturalists, from among the San-related hunter-gatherers, a six-year-old boy may have died from schistosomiasis. In Botswana, a man, who dates to 1400 BP, may have also carried the Duffy negative variant for resistance against malaria.

The genomes of Africans commonly found to undergo adaptation are regulatory DNA, and many cases of adaptation found among Africans relate to diet, physiology, and evolutionary pressures from pathogens. Throughout Sub-Saharan Africa, genetic adaptation (e.g., rs334 mutation, Duffy blood group, increased rates of G6PD deficiency, sickle cell disease) to malaria has been found among Sub-Saharan Africans, which may have initially developed in 7300 BP. Sub-Saharan Africans have more than 90% of the Duffy-null genotype. In the Kalahari Desert region of Africa, various possible genetic adaptations (e.g., adiponectin, body mass index, metabolism) have been found among the ǂKhomani peopleSub-Saharan Africans have more than 90% of the Duffy-null genotype. In South Africa, genetic adaptation (e.g., rs28647531 on chromosome 4q22) and strong susceptibility to tuberculosis has been found among Coloureds.

Recent African origin of modern humans

Between 500,000 BP and 300,000 BP, anatomically modern humans may have emerged in Africa. As Africans (e.g., Y-Chromosomal Adam, Mitochondrial Eve) have migrated from their places of origin in Africa to other locations in Africa, and as the time of divergence for East African, Central African, and West African lineages are similar to the time of divergence for the Southern African lineage, there is insufficient evidence to identify a specific region for the origin of humans in Africa. In 100,000 BP, anatomically modern humans migrated from Africa into Eurasia. Subsequently, tens of thousands of years after, the ancestors of all present-day Eurasians migrated from Africa into Eurasia and eventually became admixed with Denisovans and Neanderthals.

Archaeological and fossil evidence provide support for the African origin of homo sapiens and behavioral modernity. Models reflecting a pan-African origin (multiple locations of origin within Africa) and evolution of modern humans have been developed. As the idea of "modern" has become increasingly problematized, research has "begun to disentangle what is meant by "modern" genetic ancestry, skeletal morphology, and behavior, recognizing these are unlikely to form a single package."

In comparison to the non-African genome, the African genome features a ~25% greater number of polymorphisms, or 3 to 5 times as many, and genetic variants that are rare outside of Africa are found to occur at an abundant rate within Africa. Most of the genetic diversity found among non-Africans is found to be, at large, a subset of genetic diversity found among Africans. The genomes of Africans commonly found to undergo adaptation are regulatory DNA, and many cases of adaptation found among Africans relate to diet, physiology, and evolutionary pressures from pathogens. Throughout Sub-Saharan Africa, genetic adaptation (e.g., rs334 mutation, Duffy blood group, increased rates of G6PD deficiency, sickle cell disease) to malaria has been found among Sub-Saharan Africans, which may have initially developed in 7300 BP. Throughout Africa, various genetic adaptations (e.g., apolipoprotein L1 (APOL1): G1 and G2 haplotype resistance to trypanosomiasis and increased risk of kidney disease; human leukocyte antigen (HLA) genes; major histocompatibility complex (MHC)) to HIV-1, smallpox, trypanosomiasis (African sleeping sickness), and tuberculosis has been found among AfricansBiomedical tests for specific genetic variants (e.g., rs1799853 in the CYP2C9 gene), which have been approved by the U.S. Food and Drug Administration and are intended to indicate correct prescription of warfarin, has been found to be increasingly irrelevant to Africans as the variants are rare in Africa. As frequency rate factors into considering and deciding variant pathogenicity and generalizable polygenic scores, modern clinical classifications of genetic variant pathogenicity are found to be inadequate due to a lack of genetic diversity in biomedical studies. Fan et al (2023) recently found ~5.3 million unique genetic variants in 180 African hunter-gatherer populations, and among existing classifications for variants determined to likely be "pathogenic", ~29% (44/154) of these "pathogenic" classified variants were found to occur frequently among the African hunter-gatherers.

Haplotype

From Wikipedia, the free encyclopedia https://en.wikipedia.org/wiki/Haplotype ...