Southern and eastern African populations that speak non-Bantu languages with click consonants are known to harbour some of the most ancient genetic lineages in humans, but their relationships are poorly understood. Here, we report data from 23 populations analysed at over half a million single-nucleotide polymorphisms, using a genome-wide array designed for studying human history. The southern African Khoisan fall into two genetic groups, loosely corresponding to the northwestern and southeastern Kalahari, which we show separated within the last 30,000 years. We find that all individuals derive at least a few percent of their genomes from admixture with non-Khoisan populations that began ∼1,200 years ago. In addition, the East African Hadza and Sandawe derive a fraction of their ancestry from admixture with a population related to the Khoisan, supporting the hypothesis of an ancient link between southern and eastern Africa.

Preface and acknowledgments v List of maps, tables, and figures xiv Abbreviations and conventions xviii Representation of linguistic examples xviii Abbreviations xix Other symbols and fonts xxi 1 The framework, aim, and data of this study 1 1.1 Topic and research questions 1 1.2 Theoretical preliminaries 3 1.2.1 Reported discourse and its categories 4 1.2.2 Quotative indexes as formal signals of reported discourse 1.3 Methodological preliminaries 1.3.1 Language sample, sources, and data corpus 1.3.2 The analysis of quotative indexes 1.3.3 Elements of quotative indexes used with other functions 1.4 An overview of reported-discourse constructions in the corpus 1.4.1 Non-direct reported discourse 1.4.2 Free direct reported discourse 2 The internal structure of quotative indexes 2.1 The predicative assertion and predication operators 2.1.1 Predicative vs. non-predicative quotative indexes 2.1.2 Deviation from canonical predicativity ("marked predicativity") 2.1.3 Marked lexical features of predicative items 2.1.

The Khoisan populations of southern Africa are known to harbor some of the deepest-rooting lineages of human mtDNA; however, their relationships are as yet poorly understood. Here, we report the results of analyses of complete mtDNA genome sequences from nearly 700 individuals representing 26 populations of southern Africa who speak diverse Khoisan and Bantu languages. Our data reveal a multilayered history of the indigenous populations of southern Africa, who are likely to be the result of admixture of different genetic substrates, such as resident forager populations and pre-Bantu pastoralists from East Africa. We find high levels of genetic differentiation of the Khoisan populations, which can be explained by the effect of drift together with a partial uxorilocal/multilocal residence pattern. Furthermore, there is evidence of extensive contact, not only between geographically proximate groups, but also across wider areas. The results of this contact, which may have played a role in the diffusion of common cultural and linguistic features, are especially evident in the Khoisan populations of the central Kalahari.

Clicks are often considered an exotic feature of languages, and the fact that certain African “Khoisan” groups share the use of clicks as consonants and exhibit deep genetic divergences has been argued to indicate that clicks trace back to an early common ancestral language ( Knight et al. 2003 ). Here, we review the linguistic evidence concerning the use of click sounds in languages and the genetic evidence concerning the relationships of African click-speaking groups. The linguistic evidence suggests that genealogical inheritance and contact-induced transmission are equally relevant for the distribution of clicks in African languages. The genetic evidence indicates that there has been substantial genetic drift in some groups, obscuring their genetic relationships. Overall, the presence of clicks in human languages may in fact not trace back to the dawn of human language, but instead reflect a much later episode in the diversification of human speech.

Predication focus — a category where the predicate or a part thereof constitutes (or is part of) the sentence focus — is frequently encoded across the Bantu family by inflectional or morphosyntactic means. This phenomenon is associated with another observation which is rather unexpected at first glance. There often exists a formal parallel between marking devices of predication focus on the one hand and of present progressive on the other. This is valid across Bantu for a number of different morphological or syntactic forms. Some cases even suggest that this “isomorphism” can result from a directional grammaticalization change from predication focus toward progressive. As the formal and historical relation between the two categories cannot be viewed as accidental, an explanation is called for. Although progressivity and focus pertain to different functional domains, their relationship can indeed be motivated. The present proposal elaborates the previous hypothesis by Hyman and Watters (1984) that the progressive is an inherently focused verb category. Thus, the paper throws light in particular on the pragmatic import of progressives beyond their semantic aspect of time marking.

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Objectives: We investigated the genetic history of southern African populations with a special focus on their paternal history. We reexamined previous claims that the Y-chromosome haplogroup E1b1b (E-M293) was brought to southern Africa by pastoralists from eastern Africa, and investigated patterns of sex-biased gene flow in southern Africa. Materials and methods:We analyzed previously published complete mtDNA genome sequences and 900 kb of NRY sequences from 23 populations from Namibia, Botswana, and Zambia, as well as haplogroup frequencies from a large sample of southern African populations and 23 newly genotyped Y-linked STR loci for samples assigned to haplogroup E1b1b. Results:Our results support an eastern African origin for Y-chromosome haplogroup E1b1b (E-M293); however, its current distribution in southern Africa is not strongly associated with pastoralism, suggesting more complex demographic events and/or changes in subsistence practices in this region. The Bantu expansion in southern Africa had a notable genetic impact and was probably a rapid, male-dominated expansion. Our finding of a significant increase in the intensity of the sex-biased gene flow from north to south may reflect changes in the social dynamics between Khoisan and Bantu groups over time.Conclusions: Our study shows that the population history of southern Africa has been complex, with different immigrating groups mixing to different degrees with the autochthonous populations. The Bantu expansion led to heavily sex-biased admixture as a result of interactions between Khoisan females and Bantu males, with a geographic gradient which may reflect changes in the social dynamics between Khoisan and Bantu groups over time.

Bagirmi 2 , and Moru-Mangbetu. The two easternmost families of Niger-Congo, Benue-Congo and Adamawa-Ubangi, as well as the two Central Sudanic families, Bongo-Bagirmi and Moru-Mangbetu, will be shown to hold a particularly prominent position in the identified core group and form again a compact geographical block. Some lineages, which in geographical terms are all peripheral but still adjacent to the core, display an ambiguous behavior regarding linguistic commonalities with this area. They are Atlantic, Dogon, Songhai, Chadic, Ijoid, Narrow Bantu, and Nilotic. From the above remarks it will be clear that genealogical language groups to be considered in this paper are usually low-level units, called here "families", not the four super-groups proposed by Greenberg (1963). The discussion of why I take such smaller genealogical units as the reference of continental sampling will be postponed to Section 4.2. Suffice it to say here that this approach has the advantage that a greater variety of languages will have to be included and no relevant genealogical group for which data are available is unduly omitted. Clearly, if this breakdown would be transferred into a genealogical classification this would be a far more splitting one. The present schema, which does not refer to groups like Khoisan, Nilo-Saharan, and Niger-Kordofanian 3 , should, however, not be viewed as an alternative classification proposal, which would be a major endeavor in its own right. Low-level sampling is rather warranted by the particular topic of this paper which must consider the possibility that certain types of commonalities involving genealogical entities that are not yet based on solid evidence may well have an explanation other than common inheritance, inter alia, one in terms of areal contact. 2 Francophone specialists call this family "Sara-Bongo-Bagirmi". 3 A genealogical unit "Narrow Niger-Congo", which is on the higher-order level of a stock, rather than a family, will be assumed here, though, and includes Kru, Gur, Kwa, Benue-Congo, and Adamawa-Ubangi. This is quite comparable to a concept first developed by Stewart (1976) and Bennett and Sterk (1977) called there "Volta-Congo" and "Central Niger-Congo", respectively. While nothing in the discussion hinges on this choice, I will stick to Greenberg's usage of "Niger-Kordofanian", "Niger-Congo", etc. The main reason is that it is unclear to me which of the Post-Greenbergian classificatory and terminological proposals will eventually prevail.