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The Late Quaternary Hominins of Africa: The Skeletal Evidence from MIS 6-2

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Africa from MIS 6-2

Part of the book series: Vertebrate Paleobiology and Paleoanthropology ((VERT))

Abstract

The late Quaternary African hominin fossil record provides a tantalizing glimpse into considerable temporal and geographic morphological diversity within the genus Homo. A total of 50 sites that can be constrained from MIS 6-2 have yielded specimens ranging from isolated teeth to nearly complete skeletons. However, only a dozen or so provide particularly informative or interesting evidence spanning this period of nearly 200 kyr. In addition to the rather paltry nature of the record, one of the seemingly more intractable problems that bedevil its interpretation is the nature of the chronometric record for many of the sites. The Late Pleistocene terrestrial climatic record for Africa is also rather patchy, making continent-wide generalizations difficult. Attempts to link large-scale environmental perturbations in Africa to patterns of human evolution and behavior are even more problematic. Although the African fossil (and archaeological) record is most often viewed from the perspective of a single lineage culminating in the appearance of Homo sapiens and thence modern humans, the degree of morphological diversity evident even in this meager assemblage can be rather striking. Some of this diversity may be related to geographic and/or temporal differences, but in other instances, there are noticeable differences among remains that are contemporaneous, or at least penecontemporaneous. The Late Pleistocene African hominin fossil record, despite its manifestly incomplete nature, finds consistency with an impressive array of genetic evidence that points to an African origin for our species, and it also has consilience with genetic data that indicate a coalescence of lineages to the common ancestor of Homo sapiens at around the beginning of MIS 6. Although multiple lines of genetic evidence indicate a deep separation of lineages, with the ancestors of the southern African Khoesan diverging early on from that which gave rise to all other groups, there is a notable paucity of human remains that predate MIS 2 that exhibit strong phenetic resemblance to recent African populations. A number of the human dental samples from Late Pleistocene South African sites possess morphological variants that characterize the teeth of the recent inhabitants of sub-Saharan Africa, but these similarities do not necessarily signify a close evolutionary relationship with any of these populations because they appear to be plesiomorphic.

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Notes

  1. 1.

    Since the mid-1980s, the lower boundary of the Pleistocene Epoch (i.e., the beginning of the Quaternary Period) has been regarded as corresponding with the base of the Calabrian stratotype at 1.81 Ma. Recently, however, the International Union of Geological Sciences has recognized the base of the Gelasian stratotype, which corresponds to the Matuyama (C2r) chronozone, or the Gauss-Matuyama boundary, as defining the Pliocene-Pleistocene boundary at 2.588 Ma (Gibbard and Head 2009). This change is significant for discussions of hominin paleontology. Pending the outcome of appeals to this ruling, I continue to regard the base of the Pleistocene at 1.81 Ma.

  2. 2.

    The ages for MIS 5c employed by Shackleton (1982) and Deacon et al. (1988) correspond to the SPECMAP dates recorded in Imbrie et al. (1984).

  3. 3.

    The best-represented element from the lower part of the SAS member is the mandibular corpus (n = 4). Of these four, the KRM 41815 jaw derives from the deposits in cave 1B (Layer 10 of Singer and Wymer 1982), while the others are from separate horizons in cave 1 (KRM 16424 from Layer 14+, KRM 13400 from Layer 14, and KRM 21776 from Layer 17 of Singer and Wymer 1982).

  4. 4.

    There are some theoretical expectations and empirical observations to the effect that admixture may result in the expression of increased morphological anomalies, or of morphologies that may be intermediate in nature but with heightened levels or unusual modes of variation (Ackermann 2010). Still, we have very little expectation of how to recognize a hybrid individual (or sample) in the paleontological record. A number of specimens from sites in Europe that date to MIS 3 have been posited to represent hybrids, with most being viewed as evincing an overall modern aspect with some Neandertal features. These include the remains from Abrigo do Lagar Velho, Portugal (Duarte et al. 1999; Bayle et al. 2010), Peştera cu Oase, Romaina (Trinkaus et al. 2003; Rougier et al. 2007), Peştera Muierii, Romania (Soficaru et al. 2006), Mladeč, Czech Republic (Frayer et al. 2006; Wolpoff et al. 2006) and Cioclovina, Romania (Soficaru et al. 2007). These claims have not gone unchallenged (e.g., Tattersall and Schwartz 1999; Harvati et al. 2007). Perhaps one of the more interesting specimens in this regard is the 40–30 ka mandibular corpus from Riparo di Mezzena, Italy. This fragment has been interepreted as a Neandertal on the basis of its mtDNA, which shows “a classic Neandertal motif with the diagnostic transversion 16256 C/A” (Condemi et al. 2013: 6). However, it also displays an incipient mental trigone (chin), a feature that long has been held to be a distinction of Homo sapiens (Schwartz and Tattersall 2000).

References

  • Ackermann, R. R. (2010). Phenotypic traits of primate hybrids: Recognizing admixture in the fossil record. Evolutionary Anthropology, 19, 258–270.

    Article  Google Scholar 

  • Adams, J. M. (1997). Global land environments since the Last Interglacial. Oak Ridge National Laboratory (TN, USA). http://www.esd.ornl.gov/ern/qen/nerc.html.

  • Adkins, J., deMenocal, P., & Eshel, G. (2006). The “African Humid Period” and the record of marine upwelling from excess 230Th in ODP Hole 658C. Paleoceanography, 21, PA4203.

    Google Scholar 

  • Ahern, J. C. M., & Smith, F. H. (2004). Adolescent archaic or adult moderns? Le Moustier 1 as a model for estimating age at death of fragmentary supraorbital fossils in the modern human origins debate. Homo, 55, 1–19.

    Article  Google Scholar 

  • Aiello, L. C. (2010). Five years of Homo floresiensis. American Journal of Physical Anthropology, 142, 167–179.

    Google Scholar 

  • Aitken, M. J., & Stokes, S. (1997). Climatostratigraphy. In R. E. Taylor & M. J. Aitken (Eds.), Chronometric dating in archaeology (pp. 1–30). New York: Plenum.

    Chapter  Google Scholar 

  • Allsworth-Jones, P., Harvati, K., & Stringer, C. B. (2010). The archaeological context of the Iwo Eleru cranium from Nigeria and preliminary results of new morphometric studies. In P. Allsworth-Jones (Ed.), West African archaeology: New developments, new perspectives (BAR International Series No. 2164) (pp. 29–42). Oxford: Archaeopress.

    Google Scholar 

  • Amani, F., & Geraads, D. (1993). The Mousterian site of Jebel-Irhoud, Morocco—new data on the fauna and biochronology, and description of a new human fossil. Comptes rendus de l’Académie des sciences Série II, 316, 847–852.

    Google Scholar 

  • Ambergen, A. W., & Schaafsma, W. (1984). Interval estimates for posterior probabilities: Applications to Border Cave. In G. N. van Vark & W. W. Howells (Eds.), Multivariate statistical methods in physical anthropology (pp. 115–134). Dordrecht: Reidel.

    Chapter  Google Scholar 

  • Anderson, J. E. (1968). Late Paleolithic skeletal remains from Nubia. In F. Wendorf (Ed.), The prehistory of Nubia (2nd ed., pp. 996–1040). Dallas: Southern Methodist University Press.

    Google Scholar 

  • Andrews, J. T. (2000). Dating glacial events and correlation to global climate change. In J. S. Noller, J. M. Sowers, & W. R. Lettis (Eds.), Quaternary geology. Methods and applications (pp. 447–455). Washington, DC: American Geophysical Union.

    Chapter  Google Scholar 

  • Anthony, B. W. (1967). Excavations at Peers Cave, Fish Hoek, South Africa. Paleoecologia Africana, 2, 58–59.

    Google Scholar 

  • Antoine, D. M., Zazzo, A., & Friedman, R. (2013). Revisiting Jebel Sahaba: New apatite radiocarbon dates for one of the Nile valley’s earliest cemeteries. American Journal of Physical Anthropology, 150(S56), 68.

    Google Scholar 

  • Aouraghie, H., & Abbassi, M. (2002). Les rongeurs du site Atérien d’El Harhoura 1 (Témara, Maroc). Quaternaire, 13, 125–136.

    Article  Google Scholar 

  • Arambourg, C., Boule, M., Vallois, H., & Verneau, R. (1934). Les Grottes Paléolithiques des Béni-Ségoual (Algérie) (Archives de l’IPH No. 13). Paris: Masson.

    Google Scholar 

  • Armelagos, G. (1964). A fossilized mandible from near Wadi Halfa, Sudan. Man, 64, 12–13.

    Article  Google Scholar 

  • Armelagos, G., Van Gerven, D., Martin, D., & Huss-Ashmore, R. (1984). Effects of nutritional change on the skeletal biology of northeast African (Sudanese Nubian) populations. In J. D. Clark & S. Brandt (Eds.), From hunters to farmers: The causes and consequences of food production in Africa (pp. 132–146). Berkeley: University of California Press.

    Google Scholar 

  • Armitage, S. J., Drake, N. A., Stokes, S., El-Hawat, A., Salem, M., White, K., et al. (2007). Multiple phases of North African humidity recorded in lacustrine sediments from the Fazzan Basin, Libyan Sahara. Quaternary Geochronology, 2, 181–186.

    Article  Google Scholar 

  • Assefa, Z. (2006). Faunal remains from Porc-Épic: Paleoecological and zooarchaeological investigations from a Middle Stone Age site in southeastern Ethiopia. Journal of Human Evolution, 51, 50–75.

    Article  Google Scholar 

  • Aubert, M., Pike, A. W. G., Stringer, C., Bartsiokas, A., Kinsley, L., Eggins, S., et al. (2012). Confirmation of a late Middle Pleistocene age for the Omo Kibish 1 cranium by direct uranium-series dating. Journal of Human Evolution, 63, 704–710.

    Article  Google Scholar 

  • Avery, D. M. (1982). Micromammals as paleoenvironmental indicators and an interpretation of the Late Quaternary in the southern Cape Province, South Africa. Annals of the South African Museum, 85, 183–377.

    Google Scholar 

  • Avery, G., Cruz-Uribe, K., Goldberg, P., Grine, F. E., Klein, R., Lenardi, M. J., et al. (1997). The 1992–93 excavations at the Die Kelders Middle and Later Stone Age cave site, South Africa. Journal of Field Archaeology, 24, 263–291.

    Google Scholar 

  • Avery, G., Halkett, D., Orton, J., Steele, T., Tusenius, M., & Klein, R. (2008). The Ysterfontein 1 Middle Stone Age rock shelter and the evolution of coastal foraging. South African Archaeological Society Goodwin Series, 10, 66–89.

    Google Scholar 

  • Backwell, L., d’Errico, F., & Wadley, L. (2008). Middle Stone Age bone tools from the Howiesons Poort layers, Sibudu Cave, South Africa. Journal of Archaeological Science, 35, 1566–1580.

    Article  Google Scholar 

  • Backwell, L. R., Parkinson, A. H., Roberts, E. M., d’Errico, F., & Huchet, J. B. (2012). Criteria for identifying bone modification by termites in the fossil record. Paleogeography, Paleoclimatology, Paleoecology, 337, 72–87.

    Article  Google Scholar 

  • Balter, M. (2011). Was North Africa the launch pad for modern human migration? Science, 331, 20–23.

    Article  Google Scholar 

  • Balzeau, A., & Badawi-Fayad, J. (2005). La morphologie externe et interne de la région supra-orbitaire est-elle corrélée à des contraintes biomécaniques? Analyses structurales des populations d’Homo sapiens d’Afalou Bou Rhummel (Algérie) et de Taforalt (Maroc). Bulletins et Mémoires de la Société d’Anthropologie de Paris, 17, 185–197.

    Google Scholar 

  • Barham, L. (1995). The Mumbwa Caves Project, Zambia, 1993-94. Nyame Akuma, 43, 66–72.

    Google Scholar 

  • Barham, L. (2000). The Mumbwa Caves 1993–1996. In: L. Barham (Ed.), The Middle Stone Age of Zambia, South Central Africa (pp. 11–49). Bristol: Western Academic & Specialist Press.

    Google Scholar 

  • Barham, L. (2002). Backed tools in Middle Pleistocene central Africa and their evolutionary significance. Journal of Human Evolution, 43, 585–603.

    Article  Google Scholar 

  • Barham, L., & Debenham, N. (2000). Mumbwa Caves chronology. In L. Barham (Ed.), The Middle Stone Age of Zambia, south central Africa (pp. 43–49). Bristol, UK: Western Academic & Specialist Press.

    Google Scholar 

  • Barham, L., Simms, M., Gilmour, M., & Debenham, N. (2000). Twin Rivers, excavation and behavioural record. In L. Barham (Ed.), The Middle Stone Age of Zambia, south central Africa (pp. 165–216). Bristol, UK: Western Academic & Specialist Press.

    Google Scholar 

  • Barkai, R., Gopher, A., Lauritzen, S. E., & Frumkin, A. (2003). Uranium series dates from Qesem Cave, Israel, and the end of the Lower Paleolithic. Nature, 423, 977–979.

    Article  Google Scholar 

  • Barkai, R., Gopher, A., & Shimelmitz, R. (2005). Middle Pleistocene blade production in the Levant: An Amudian assemblage from Qesem Cave, Israel. Eurasian Prehistory, 3, 39–74.

    Google Scholar 

  • Barker, G., Hunt, C., & Reynolds, T. (2007). The Haua Fteah, Cyrenaica (Northeast Libya): Renewed investigations of the cave and its landscape, 2007. Libyan Studies, 38, 1–22.

    Article  Google Scholar 

  • Barker, G., Antoniadou, A., Armitage, S., Brooks, I., Candy, I., Connell, K., et al. (2010). The Cyrenaican Prehistory Project 2010: The fourth season of investigations of the Haua Fteah cave and its landscape, and further results from the 2007-2008 fieldwork. Libyan Studies, 41, 63–88.

    Article  Google Scholar 

  • Barker, G., Bennett, P., Farr, L., Hill, E., Hunt, C., Lucarini, G., et al. (2012). The Cyrenaican Prehistory Project 2012: The fifth season of investigations of the Haua Fteah cave. Libyan Studies, 43, 115–136.

    Article  Google Scholar 

  • Barker, P., & Gasse, F. (2003). New evidence for a reduced water balance in East Africa during the Last Glacial Maximum: Implication for model-data comparison. Quaternary Science Reviews, 22, 823–837.

    Article  Google Scholar 

  • Bar-Matthews, M., Marean, C. W., Jacobs, Z., Karkanas, P., Fisher, E. C., Herries, A. I. R., et al. (2010). A high resolution and continuous isotopic speleothem record of paleoclimate and paleoenvironment from 90 to 53 ka from Pinnacle Point on the south coast of South Africa. Quaternary Science Reviews, 29, 2131–2145.

    Article  Google Scholar 

  • Barton, R. N. E., Bouzouggar, J., Collcutt, S. N., Gale, R., Higham, T. F. G., Humphrey, L. T., et al. (2005). The late Upper Paleolithic occupation of the Moroccan northwest Maghreb during the Last Glacial Maximum. African Archaeological Review, 22, 77–100.

    Article  Google Scholar 

  • Barton, R. N. E., Bouzouggar, A., Bronk-Ramsey, C., Collcutt, S. N., Higham, T. F. G., Humphrey, L. T., et al. (2007). Abrupt climatic change and chronology of the Upper Paleolithic in northern and eastern Morocco. In O. Bar-Yosef, P. Mellars, C. Stringer, & K. Boyle (Eds.), Rethinking the human revolution: New behavioural and biological perspectives on the origins and dispersal of modern humans (pp. 177–186). Cambridge: McDonald Institute for Archaeological Research.

    Google Scholar 

  • Barton, N., Bouzouggar, A., Humphrey, L., Berridge, P., Collcutt, S., Gale, R., et al. (2008). Human burial evidence from Hattab II Cave and the question of continuity in Late Pleistocene – Holocene mortuary practices in northwest Africa. Cambridge Archaeological Journal, 18, 195–214.

    Article  Google Scholar 

  • Barton, R. N. E., Bouzouggar, A., Collcutt, S. N., Schwenninger, J. L., & Clark-Balzan, L. (2009). OSL dating of the Aterian levels at Dar es-Soltan I (Rabat, Morocco) and implications for the dispersal of modern Homo sapiens. Quaternary Science Reviews, 28, 1914–1931.

    Article  Google Scholar 

  • Barton, R. N. E., Bouzouggar, A., Hogue, J. T., Lee, S., Collcutt, S. N., & Ditchfield, P. (2013). Origins of the Iberomaurusian in NW Africa: New AMS radiocarbon dating of the Middle and Later Stone Age deposits at Taforalt Cave, Morocco. Journal of Human Evolution, 65, 266–281.

    Article  Google Scholar 

  • Bassinot, F. C., Labeyrie, L. D., Vincent, E., Quidelleur, X., Shackleton, N. J., & Lancelot, Y. (1994). The astronomical theory of climate and the age of the Brunhes-Matuyama magnetic reversal. Earth and Planetary Science Letters, 126, 91–108.

    Article  Google Scholar 

  • Bayle, P., Macchiarelli, R., Trinkaus, E., Duarte, C., Mazurier, A., & Zilhão, J. (2010). Dental maturational sequence and dental tissue proportions in the early Upper Paleolithic child from Abrigo do Lagar Velho, Portugal. Proceedings of the National Academy of Sciences of the United States of America, 107, 1338–1342.

    Article  Google Scholar 

  • Beaumont, P. B. (1980). On the age of Border Cave hominids 1-5. Paleontologia Africana, 23, 21–33.

    Google Scholar 

  • Beaumont, P. B., de Villiers, H., & Vogel, J. C. (1978). Modern man in sub-Saharan Africa prior to 49,000 years B.P.: A review and evaluation with particular reference to Border Cave. South African Journal of Science, 74, 409–419.

    Google Scholar 

  • Behar, D. M., & The Genographic Consortium. (2008). The dawn of human matrilineal diversity. American Journal of Human Genetics, 82, 1130–1140.

    Article  Google Scholar 

  • Behrensmeyer, A. K. (2006). Climate change and human evolution. Science, 311, 476–478.

    Article  Google Scholar 

  • Ben-Ncer, A. (2004). Etude de la sépulture Ibéromaurusienne 1 d’Ifri n’Baroud (Rif oriental, Maroc). Anthropo, 7, 177–185.

    Google Scholar 

  • Biittner, K. (2011). Characterization of Middle and Later Stone Age lithic artifacts from two rockshelter sites in Iringa Region, southern Tanzania. Ph.D. Dissertation, University of Alberta.

    Google Scholar 

  • Biittner, K. M., Bushozi, P. M., & Willoughby, P. R. (2007). The Middle and Later Stone Age of the Iringa Region, southern Tanzania: An introduction. Nyame Akuma, 68, 62–73.

    Google Scholar 

  • Bird, M. I., Fifield, L. K., Santos, G. M., Beaumont, P. B., Zhou, Y., di Tada, M. L., et al. (2003). Radiocarbon dating from 40 to 60 ka BP at Border Cave, South Africa. Quaternary Science Reviews, 22, 943–947.

    Article  Google Scholar 

  • Blegen, N., Tryon, C. A., Faith, J. T., Peppe, D. J., Beverly, E. J., Li, B., & Jacobs, Z. (2015). Distal tephras of the eastern Lake Victoria Basin, equatorial East Africa: Correlations, chronology, and a context for early modern humans. Quaternary Science Reviews, 122, 89–111.

    Google Scholar 

  • Blome, M. W., Cohen, A. S., Tryon, C. A., Brooks, A. S., & Russell, J. (2012). The environmental context for the origins of modern human diversity: A synthesis of regional variability in African climate 150,000-30,000 years ago. Journal of Human Evolution, 62, 563–592.

    Article  Google Scholar 

  • Boaz, N. T., Pavlakis, P., & Brooks, A. S. (1990). Late Pleistocene-Holocene human remains from the Upper Semliki, Zaire. In N. T. Boaz (Ed.), Evolution of environments and hominidae in the African Western Rift Valley (Memoir No. 1) (pp. 273–299). Martinsville, VA: Virginia Museum of Natural History.

    Google Scholar 

  • Bond, G., Broecker, W., Johnsen, S., McManus, J., Labeyrie, L., Jouzel, J., et al. (1993). Correlations between climate records from North Atlantic sediments and Greenland ice. Nature, 365, 143–147.

    Article  Google Scholar 

  • Bouchneb, L., & Crevecoeur, I. (2009). The inner ear of Nazlet Khater 2 (Upper Paleolithic, Egypt). Journal of Human Evolution, 56, 257–262.

    Article  Google Scholar 

  • Bousman, B. C. (2005). Coping with risk: Later Stone Age technological strategies at Blydefontein Rock Shelter, South Africa. Journal of Anthropological Archaeology, 24, 193–226.

    Article  Google Scholar 

  • Bouzouggar, A., Barton, N., Vanhaeren, M., d’Errico, F., Collcutt, S., Higham, T., et al. (2007). 82,000 year-old shell beads from North Africa and implications for the origins of modern human behavior. Proceedings of the National Academy of Sciences of the United States of America, 104, 9964–9969.

    Article  Google Scholar 

  • Bouzouggar, A., Barton, R. N. E., Blockley, S., Bronk-Ramsey, C., Collcutt, S. N., Gale, R., et al. (2008). Reevaluating the age of the Iberomaurusian in Morocco. African Archaeological Review, 25, 3–19.

    Article  Google Scholar 

  • Bowen, B. E., & Vondra, C. F. (1973). Stratigraphical relationships of the Plio-Pleistocene deposits, East Rudolf, Kenya. Nature, 242, 391–393.

    Article  Google Scholar 

  • Bräuer, G. (1984a). The Afro-European sapiens hypothesis and hominid evolution in East Asia during the late Middle Pleistocene and Upper Pleistocene. Courier Forschungsinstitut Senckenberg, 69, 145–165.

    Google Scholar 

  • Bräuer, G. (1984b). A craniological approach to the origin of anatomically modern Homo sapiens in Africa and implications for the appearance of modern Europeans. In F. H. Smith & F. Spencer (Eds.), The origins of modern humans: A world survey of the fossil evidence (pp. 327–410). New York: Alan R. Liss.

    Google Scholar 

  • Bräuer, G. (1989a). The evolution of modern humans: A comparison of the African and non-African evidence. In P. Mellars & C. Stringer (Eds.), The human revolution (pp. 123–154). Edinburgh: Edinburgh University Press.

    Google Scholar 

  • Bräuer, G. (1989b). The ES-11693 hominid from West Turkana and Homo sapiens evolution in East Africa. In G. Giacobini (Ed.), Hominidae (pp. 241–245). Milan: Jaca Books.

    Google Scholar 

  • Bräuer, G. (1992). Africa’s place in the evolution of Homo sapiens. In G. Bräuer & F. H. Smith (Eds.), Continuity or replacement? Controversies in Homo sapiens evolution (pp. 83–98). Rotterdam: Balkema.

    Google Scholar 

  • Bräuer, G. (2001). The KNM-ER 3884 hominid and the emergence of modern human anatomy in Africa. In P. V. Tobias, M. A. Rath, J. Moggi-Cecchi, & G. A. Doyle (Eds.), Humanity from African naissance to coming millennia (pp. 191–197). Firenze: Firenza University Press.

    Google Scholar 

  • Bräuer, G. (2008). The origin of modern anatomy: By speciation or intraspecific evolution? Evolutionary Anthropology, 17, 22–37.

    Article  Google Scholar 

  • Bräuer, G., & Leakey, R. E. (1986a). A new archaic Homo sapiens cranium from Eliye Springs, West Turkana, Kenya. Zeitschrift für Morphologie und Anthropologie, 76, 245–252.

    Google Scholar 

  • Bräuer, G., & Leakey, R. E. (1986b). The ES-11693 cranium from Eliye Springs, West Turkana, Kenya. Journal of Human Evolution, 15, 289–312.

    Article  Google Scholar 

  • Bräuer, G., & Mabulla, Z. P. (1996). New hominid fossil from Lake Eyasi, Tanzania. Anthropologie, 34, 47–53.

    Google Scholar 

  • Bräuer, G., & Mehlman, M. J. (1988). Hominid molars from a Middle Stone Age level at the Mumba Rock Shelter, Tanzania. American Journal of Physical Anthropology, 75, 69–76.

    Article  Google Scholar 

  • Bräuer, G., & Rimbach, K. (1990). Late archaic and modern Homo sapiens from Europe, Africa and southwest Asia: Craniometric comparisons and phylogenetic implications. Journal of Human Evolution, 19, 789–807.

    Article  Google Scholar 

  • Bräuer, G., & Rösing, F. W. (1989). Human biological history in southern Africa. Rassengeschichte der Menschheit, 13, 6–137.

    Google Scholar 

  • Bräuer, G., & Singer, R. (1996). The Klasies zygomatic bone: Archaic or modern? Journal of Human Evolution, 30, 161–165.

    Article  Google Scholar 

  • Bräuer, G., Deacon, H. J., & Zipfel, F. (1992a). Comment on the new maxillary finds from Klasies River, South Africa. Journal of Human Evolution, 23, 419–422.

    Article  Google Scholar 

  • Bräuer, G., Leakey, R. E. F., & Mbua, E. (1992b). A first report on the ER-3884 cranial remains from Ileret/East Turkana, Kenya. In G. Bräuer & F. H. Smith (Eds.), Continuity or replacement? Controversies in Homo sapiens evolution (pp. 111–120). Rotterdam: Balkema.

    Google Scholar 

  • Bräuer, G., Yokoyama, Y., Falguères, C., & Mbua, E. (1997). Modern human origins backdated. Nature, 386, 337.

    Article  Google Scholar 

  • Bräuer, G., Groden, C., Delling, G., Kupczik, K., Mbua, E., & Schultz, M. (2003). Pathological alterations in the archaic Homo sapiens cranium from Eliye Springs, Kenya. American Journal of Physical Anthropology, 120, 200–204.

    Google Scholar 

  • Bräuer, G., Groden, C., Gröning, F., Kroll, A., Kupczik, K., Mbua, E., et al. (2004). Virtual study of the endocranial morphology of the matrix-filled cranium from Eliye Springs, Kenya. Anatomicasl Record, 276A, 113–133.

    Article  Google Scholar 

  • Breuil, A. H., Teilhard de Chardin, P., & Wernert, P. (1951). Le Paléolithique du Harar. L’ Anthropologie, 55, 219–230.

    Google Scholar 

  • Briggs, L. C. (1955). The Stone Age races of northwest Africa (Bulletin of the American School of Prehistoric Research No. 18). Cambridge, MA: Peabody Museum.

    Google Scholar 

  • Brooks, A. S. (2008). Overview of the sequence, environments and dating at Ishango and Katanda. In D. Huylebrouck (Ed.), Ishango, 22000 and 50 years later: The cradle of mathematics? (pp. 55–66). Brussels: Koninklijke Vlaamse Academie van Belgie voor Wetenschappen en Kunsten.

    Google Scholar 

  • Brooks, A. S., & Robertshaw, P. (1990). The Glacial Maximum in tropical Africa: 22,000–12,000 B.P. In C. Gamble & O. Soffer (Eds.), The World at 18 000 BP: Low latitudes, Volume 2 (pp 121–169). London: Unwin Hyman.

    Google Scholar 

  • Brooks, A. S., & Smith, C. C. (1987). Ishango revisited: New age determinations and cultural interpretations. African Archaeological Review, 5, 65–78.

    Article  Google Scholar 

  • Brooks, A. S., Smith, C. C., & Boaz, N. T. (1991). New human remains from Ishango, Zaire, in relation to Later Pleistocene human evolution. American Journal of Physical Anthropology, S12, 54–55.

    Google Scholar 

  • Brooks, A. S., Helgren, D. M., Cramer, J. S., Franklin, A., Hornyak, W., Keating, J. M., et al. (1995). Dating and context of three Middle Stone Age sites with bone points in the Upper Semliki Valley, Zaire. Science, 268, 548–553.

    Article  Google Scholar 

  • Broom, R. (1918). The evidence afforded by the Boskop skull of a new species of primitive man (Homo capensis). Anthropological Papers of the American Museum of Natural History, 23, 63–79.

    Google Scholar 

  • Broom, R. (1923). A contribution to the craniology of the yellow-skinned races of South Africa. Journal of the Royal Anthropological Institute, 53, 132–149.

    Google Scholar 

  • Broom, R. (1929a). The Transvaal fossil human skeleton. Nature, 123, 415–416.

    Article  Google Scholar 

  • Broom, R. (1929b). Australoid element in the Koranas. Nature, 124, 507.

    Article  Google Scholar 

  • Brothwell, D. R. (1963). Evidence of early population change in Central and southern Africa: Doubts and problems. Man, 63, 101–104.

    Article  Google Scholar 

  • Brothwell, D., & Shaw, T. (1971). A Late Upper Pleistocene proto-West African Negro from Nigeria. Man, 2, 221–227.

    Article  Google Scholar 

  • Brown, F. H., & Feibel, C. S. (1986). Revision of lithostratigraphic nomenclature in the Koobi Fora region, Kenya. Journal of the Geological Society London, 143, 297–310.

    Article  Google Scholar 

  • Brown, F. H., & Fuller, C. (2008). Stratigraphy and tephra of the Kibish Formation, southwestern Ethiopia. Journal of Human Evolution, 55, 366–403.

    Article  Google Scholar 

  • Brown, F. H., McDougall, I., & Fleagle, J. G. (2012). Correlation of the KHS Tuff of the Kibish Formation to volcanic ash layers at other sites, and the age of early Homo sapiens (Omo I and Omo II). Journal of Human Evolution, 63, 577–585.

    Article  Google Scholar 

  • Brown, K. S., Marean, C. W., Herries, A. I. R., Jacobs, Z., Tribolo, C., Braun, D., et al. (2009). Fire as an engineering tool of early modern humans. Science, 325, 859–862.

    Article  Google Scholar 

  • Brown, K. S., Marean, C. W., Jacobs, Z., Schoville, B. J., Oestmo, S., Fisher, E. C., et al. (2012). An early and enduring advanced technology originating 71,000 years ago in South Africa. Nature, 491, 590–593.

    Article  Google Scholar 

  • Brown, P., Sutikna, T., Morwood, M. J., Soejono, R. P., Jatmiko Saptomo, E. W., et al. (2004). A new small-bodied hominin from the Late Pleistocene of Flores, Indonesia. Nature, 431, 1055–1061.

    Article  Google Scholar 

  • Bruch, A. A., Sievers, C., & Wadley, L. (2012). Quantification of climate and vegetation from southern African Middle Stone Age sites—an application using Late Pleistocene plant material from Sibudu, South Africa. Quaternary Science Reviews, 45, 7–17.

    Article  Google Scholar 

  • Bruner, E., & Manzi, G. (2002). The virtual replica of Nazlet Khater, Egypt, cranium and mandible: First results. In P. M. Vermeersch (Ed.), Paleolithic quarrying sites in Upper and Middle Egypt (pp. 337–346). Leuven: Leuven University Press.

    Google Scholar 

  • Bruzek, J., Sefcakova, A., & Cerny, V. (2004). Révision du sexe des squelettes épipaléolithiques de Taforalt et d’Afalou-bou-Rhoummel par une approche probabiliste. Antropo, 7, 195–202.

    Google Scholar 

  • Buckley, J. D., & Willis, E. H. (1969). Isotope radiocarbon measurements VII. Radiocarbon, 11, 53–105.

    Google Scholar 

  • Butler, B. H. (1974). Skeletal remains from a Late Paleolithic site near Esna. In D. Lubell (Ed.), The Fakhurian. A Late Paleolithic industry from Upper Egypt (Geological Survey of Egypt, Paper No. 58) (pp. 176–183). Cairo: The Geological Survey of Egypt.

    Google Scholar 

  • Butzer, K. W. (1969). Geological interpretation of two Pleistocene hominid sites in the Lower Omo Basin. Nature, 222, 1133–1135.

    Article  Google Scholar 

  • Butzer, K. W. (1997). Late Quaternary problems of the Egyptian Nile: Stratigraphy, environments, prehistory. Paléorient, 23, 151–173.

    Article  Google Scholar 

  • Butzer, K. W. (2004). Coastal eolian sands, paleosols, and Pleistocene geoarchaeology of the southwestern Cape, South Africa. Journal of Archaeological Science, 31, 1743–1781.

    Article  Google Scholar 

  • Butzer, K. W., & Hansen, C. L. (1968). Desert and river in Nubia: Geomorphology and prehistoric environments at the Aswan Reservoir. Madison: University of Wisconsin Press.

    Google Scholar 

  • Butzer, K. W., Brown, F. H., & Thurber, D. L. (1969). Horizontal sediments of the Lower Omo Valley: The Kibish Formation. Quaternaria, 11, 15–29.

    Google Scholar 

  • Butzer, K. W., Beaumont, P. B., & Vogel, J. C. (1978a). Lithostratigraphy of Border Cave, Kwazulu, South Africa: A Middle Stone Age sequence beginning c. 195,000 B.P. Journal of Archaeological Science, 5, 317–341.

    Article  Google Scholar 

  • Butzer, K., Stuckenrath, R., Bruzewicz, A., & Helgren, D. (1978b). Late Cenozoic paleoclimates of the Gaap Escarpment, Kalahari margin, South Africa. Quaternary Research, 10, 310–339.

    Article  Google Scholar 

  • Cambell, N. A. (1984). Some aspects of discrimination and allocation. In G. N. van Vark & W. W. Howells (Eds.), Multivariate statistical methods in physical anthropology (pp. 177–192). Dordrecht: Reidel.

    Chapter  Google Scholar 

  • Camp, A., & O’Sullivan, G. (1999). Neogene sapropels in the Mediterranean: A review. Marine Geology, 153, 11–28.

    Article  Google Scholar 

  • Camps, G. (1969). L’homme de Mechta El Arbi et sa civilisation. Contribution à l’étude des origines Guanches. Annuario de estudios atlanticos, Madrid, 15, 257–272.

    Google Scholar 

  • Cann, R. L., Stoneking, M., & Wilson, A. C. (1987). Mitochondrial DNA and human evolution. Nature, 32, 31–36.

    Article  Google Scholar 

  • Carlson, D. S. (1976). Temporal variation in prehistoric Nubian crania. American Journal of Physical Anthropology, 45, 467–484.

    Article  Google Scholar 

  • Carto, S. L., Weaver, A. J., Hetherington, R., Lam, Y., & Wiebe, E. C. (2009). Out of Africa and into an ice age: On the role of global climate change in the Late Pleistocene migration of early modern humans out of Africa. Journal of Human Evolution, 56, 139–151.

    Article  Google Scholar 

  • Castañeda, I. S., Mulitza, S., Schefuß, E., dos Santos, R. A. L., Damsté, J. S. S., & Schouten, S. (2009). Wet phases in the Sahara/Sahel region and human migration patterns in North Africa. Proceedings of the National Academy of Sciences of the United States of America, 106, 20159–20163.

    Article  Google Scholar 

  • Cerling, T. E., & Quade, J. (1993). Stable carbon and oxygen isotopes in soil carbonates. In P. K. Swart, K. C. Lohmann, J. A. McKenzie, & E. S. Savin (Eds.), Climate change in continental isotope records (pp. 217–231). Washington, DC: American Geophysical Union.

    Chapter  Google Scholar 

  • Chamla, M. C. (1970). Les hommes epipaleolithiques de la Columnata (Mémoires du C.R.A.R.E. XV). Paris: Arts et Métiers Graphiques.

    Google Scholar 

  • Chase, B. (2010). South African paleoenvironments during Marine Oxygen Isotope Stage 4: A context for the Howiesons Poort and Still Bay industries. Journal of Archaeological Science, 37, 1359–1366.

    Article  Google Scholar 

  • Chase, B. M., & Meadows, M. E. (2007). Late Quaternary dynamis of southern Africa’s winter rainfall zone. Earth Science Reviews, 84, 103–138.

    Article  Google Scholar 

  • Chavaillon, J. (1982). Position chronologique des hominidés fossile d’Ethiopie. In M. A. de Lumley (Ed.), Homo erectus et la place de l’Homme de Tautavel parmi les hominidés fossiles. Première congrès international de paléontologie humaine (pp. 766–797). UNESCO: Colloque International du Centre National de la Récherche Scientifique.

    Google Scholar 

  • Churcher, C. S. (1972). Late Pleistocene vertebrates from archaeological sites in the plain of Kom Ombo, Upper Egypt. Life Sciences Contributions of the Royal Ontario Museum, 82, 1–172.

    Google Scholar 

  • Churchill, S. E., Pearson, O. M., Grine, F. E., Trinkaus, E., & Holliday, T. W. (1996). Morphological affinities of the proximal ulna from Klasies River Main Site: Archaic or modern? Journal of Human Evolution, 31, 213–237.

    Article  Google Scholar 

  • Clark, J. D. (1942). Further excavations (1939) at Mumbwa Caves, Northern Rhodesia. Transactions of the Royal Society of South Africa, 29, 133–201.

    Article  Google Scholar 

  • Clark, J. D. (1971). Human behavioral differences in southern Africa during the later Pleistocene. American Anthropologist, 73, 1211–1236.

    Article  Google Scholar 

  • Clark, J. D., & Brown, K. S. (2001). The Twin Rivers Kopje, Zambia: Stratigraphy, fauna, and artifact assemblages from the 1954 and 1956 excavations. Journal of Archaeological Science, 28, 305–330.

    Article  Google Scholar 

  • Clark, J. D., & Williamson, K. D. (1984). A Middle Stone Age occupation site at Porc-Epic Cave, Dire Dawa (east-central Ethiopia), part I. African Archaeological Review, 2, 37–64.

    Article  Google Scholar 

  • Clark, J. D., Beyene, Y., WoldeGabriel, G., Hart, W. K., Renne, P. R., Gilbert, H., et al. (2003). Stratigraphic, chronological and behavioural contexts of Pleistocene Homo sapiens from Middle Awash, Ethiopia. Nature, 423, 747–752.

    Article  Google Scholar 

  • Clark, P. U., Dyke, A. S., Shakun, J. D., Carlson, A. E., Clark, J., Wohlfarth, B., et al. (2009). The Last Glacial Maximum. Science, 325, 710–714.

    Article  Google Scholar 

  • Close, A. E. (1988). Current research and recent radiocarbon dates from northern Africa, III. Journal of African History, 29, 145–176.

    Article  Google Scholar 

  • Close, A. E., & Wendorf, F. (1990). North Africa at 18,000 BP. In C. Gamble & O. Soffer (Eds.), The world at 18,000 BP (Vol. 2, pp. 41–57). Low Latitudes London: Unwin Hyman.

    Google Scholar 

  • Cohen, A. S., Stone, J. R., Beuning, K. R. M., Park, L. E., Reinthal, P. N., Dettman, D., et al. (2007). Ecological consequences of early Late Pleistocene megadroughts in tropical Africa. Proceedings of the National Academy of Sciences of the United States of America, 104, 16422–16427.

    Article  Google Scholar 

  • Cohen, P. (1996). Fitting a face to Ngaloba. Journal of Human Evolution, 30, 373–379.

    Article  Google Scholar 

  • Cole, J. M., Goldstein, S. L., deMenocal, P. B., Hemming, S. R., & Grousset, F. E. (2009). Contrasting compositions of Saharan dust in the eastern Atlantic Ocean during the last deglaciation and African Humid Period. Earth and Planetary Science Letters, 278, 257–266.

    Article  Google Scholar 

  • Collins, B. R., & Willoughby, P. R. (2010). The faunal analysis of Magubike and Mlambalasi, two MSA-LSA archaeological sites from Iringa District, Tanzania. Journal of Taphonomy, 8, 33–68.

    Google Scholar 

  • Compton, J. S. (2011). Pleistocene sea-level fluctuations and human evolution on the southern coastal plain of South Africa. Quaternary Science Reviews, 30, 506–527.

    Article  Google Scholar 

  • Condemi, S., Mounier, A., Giunti, P., Lari, M., Carmelli, D., & Longo, L. (2013). Possible interbreeding in late Italian Neanderthals? New data from the Mezzena Jaw (Monti Lessini, Verona, Italy). PLoS ONE, 8(3), e59781. doi:10.1371/journal.pone.0059781.

    Article  Google Scholar 

  • Cooke, H. B. S., Malan, B. D., & Wells, L. H. (1945). Fossil man in the Lebombo Mountains, South Africa: The ‘Border Cave’, Ingwavuma district, Zululand. Man, 45, 6–13.

    Article  Google Scholar 

  • Copeland, L. (2000). Yabrudian and related industries: The state of research in 1996. In A. Ronen & M. Weinstein-Evron (Eds.), Toward modern humans: Yabrudian and Micoquian, 400-50 k-years ago (BAR International Series No. 850) (pp. 97–117). Oxford: Archaeopress.

    Google Scholar 

  • Coplen, T. B. (1988). Normalization of oxygen and hydrogen isotope data. Chemical Geology: Isotope Geoscience Section, 72, 293–297.

    Google Scholar 

  • Coplen, T. B. (1996). New guidelines for reporting stable hydrogen, carbon, and oxygen isotope-ratio data. Geochimica et Cosmochimica Acta, 60, 3359–3360.

    Article  Google Scholar 

  • Cramp, A., & O’Sullivan, G. (1999). Neogene sapropels in the Mediterranean: A review. Marine Geology, 153, 11–28.

    Article  Google Scholar 

  • Cremaschi, M., Di Lernia, S., & Garcea, E. A. A. (1998). Some insights on the Aterian in the Libyan Sahara: Chronology, environment, and archeology. African Archaeological Review, 15, 261–286.

    Article  Google Scholar 

  • Crevecoeur, I. (2008a). Étude anthropologique du squelette du Paléolithique supérieur de Nazlet Khater 2 (Égypte). Apport à la compréhension de la variabilité passée des hommes modernes. Leuven: Leuven University Press.

    Google Scholar 

  • Crevecoeur, I. (2008b). Variability of Paleolithic modern humans in Africa. Future prospects of the Ishango human remain (re-)study. In D. Huylebrouck (Ed.), Ishango, 22000 and 50 years later: The cradle of mathematics? (pp. 87–97). Brussels: Koninklijke Vlaamse Academie van Belgie voor Wetenschapppen en Kunsten.

    Google Scholar 

  • Crevecoeur, I. (2012). The Upper Paleolithic human remains of Nazlet Khater 2 (Egypt) and past modern human diversity. In J. J. Hublin & S. McPherron (Eds.), Modern origins: A North African perspective (pp. 205–221). Dordrecht: Springer.

    Chapter  Google Scholar 

  • Crevecoeur, I., & Trinkaus, E. (2004). From the Nile to the Danube: A comparison of the Nazlet Khater 2 and Oase 1 early modern human mandibles. Anthropos (Brno), 42, 229–239.

    Google Scholar 

  • Crevecoeur, I., Rougier, H., Grine, F. E., & Froment, A. (2009). Modern human cranial diversity in the Late Pleistocene of Africa and Eurasia: Evidence from Nazlet Khater, Peştera cu Oase, and Hofmeyr. American Journal of Physical Anthropology, 140, 347–358.

    Article  Google Scholar 

  • Crevecoeur, I., Semal, P., Cornelissen, E., & Brooks, A. S. (2010). The Late Stone Age human remains from Ishango (Democratic Republic of Congo): Contribution to the study of the African Late Pleistocene modern human diversity. American Journal of Physical Anthropology, S50, 87.

    Google Scholar 

  • Crosta, X., & Shemesh, A. (2002). Reconciling down core anticorrelation of diatom carbon and nitrogen isotopic ratios from the Southern Ocean. Paleoceanography, 17, 10.1-10.810.1029/2000PA000565.

    Google Scholar 

  • Crowley, T. J. (1992). North Atlantic deep water cools the Southern Hemisphere. Paleoceanography, 7, 489–497.

    Article  Google Scholar 

  • Cruciani, F., Trombetta, B., Massaia, A., Destro-Bisol, G., Sellitto, D., & Scozzari, R. (2011). A revised root for the human Y chromosomal phylogenetic tree: The origin of patrilineal diversity in Africa. American Journal of Human Genetics, 88, 814–818.

    Article  Google Scholar 

  • Currat, M., & Excoffier, L. (2011). Strong reproductive isolation between humans and Neanderthals inferred from observed patterns of introgression. Proceedings of the National Academy of Sciences of the United States of America, 108, 15129–15134.

    Article  Google Scholar 

  • Damas, H. (1940). Observations sur des couches fossiliferes bordant la Semliki. Revue Zoologique et Botanique Africaine, 33, 265–272.

    Google Scholar 

  • Dansgaard, W. (1964). Stable isotopes in precipitation. Tellus, 16, 466–468.

    Article  Google Scholar 

  • Dansgaard, W., Johnsen, S. J., Clausen, H. B., Dahl-Jensen, D., Gundestrup, N. S., Hammer, C. U., et al. (1993). Evidence for general instability of past climate from a 250-ky ice-core record. Nature, 364, 218–220.

    Article  Google Scholar 

  • Dart, R. A., & del Grande, N. (1931). The ancient iron-smelting cavern at Mumbwa. Transactions of the Royal Society of South Africa, 19, 379–427.

    Google Scholar 

  • Day, M. H. (1969). Early Homo sapiens remains from the Omo River region of southwest Ethiopia. Nature, 222, 1132–1138.

    Article  Google Scholar 

  • Day, M. H., & Leakey, R. E. F. (1974). New evidence of the genus Homo from East Rudolf, Kenya (III). American Journal of Physical Anthropology, 41, 367–380.

    Article  Google Scholar 

  • Day, M. H., & Stringer, C. B. (1982). A reconsideration of the Omo Kibish remains and the erectus-sapiens transition. In M. A. de Lumley (Ed.). Homo erectus et la place de l’Homme de Tautavel parmi les hominidés fossiles. Première congrès international de paléontologie humaine (pp. 814–846). UNESCO: Colloque International du Centre National de la Récherche Scientifique.

    Google Scholar 

  • Day, M. H., & Stringer, C. B. (1991). Les restes crânes d’Omo-Kibish et leur classification à l’intérieur du genre Homo. L’Anthropologie, 95, 573–594.

    Google Scholar 

  • Day, M. H., Leakey, M. D., & Magori, C. C. (1980). A new fossil hominid skull (LH 18) from the Ngaloba Beds, Laetoli, Tanzania. Nature, 284, 55–56.

    Article  Google Scholar 

  • Deacon, H. J. (1992). Southern Africa and modern human origins. Philosophical Transactions of the Royal Society London B, 337, 177–183.

    Article  Google Scholar 

  • Deacon, H. J. (1995). Two Late Pleistocene-Holocene archaeological depositories from the southern Cape, South Africa. South African Archaeology Bulletin, 50, 121–131.

    Article  Google Scholar 

  • Deacon, H. J. (2008). The context of the 1967-8 sample of human remains from Cave 1 Klasies River Main Site. South African Archaeological Society Goodwin Series, 10, 143–149.

    Google Scholar 

  • Deacon, H. J., & Geleijnse, V. B. (1988). The stratigraphy and sedimentology of the Main Site sequence, Klasies River, South Africa. South African Archaeology Bulletin, 43, 5–14.

    Article  Google Scholar 

  • Deacon, H. J., Talma, A. S., & Vogel, J. C. (1988). Biological and cultural development of Pleistocene people in an Old World southern continent. In J. R. Prescott (Ed.), Early man in the southern Hemisphere (pp. S23–S31). Adelaide: University of Adelaide, Department of Physics and Mathematical Physics.

    Google Scholar 

  • Debénath, A. (1976). Le site de Dar-es-Soltane 2, à Rabat (Maroc). Bulletins et Mémoires de la Société d’Anthropologie de Paris, 3, 181–182.

    Article  Google Scholar 

  • Debénath, A. (1979). Découverte d’une mandibule humaine Atérienne à El Harhoura (Province de Rabat). Bulletin d’archéologie marocaine, 12, 1–2.

    Google Scholar 

  • Debénath, A. (1980). Nouveaux restes humains Atériens du Maroc. Comptes Rendus des Séances de l’Académie des Sciences (Paris) Serie D, 290, 851–852.

    Google Scholar 

  • Debénath, A. (2000). Le peuplement préhistorique du Maroc: Données récentes et problèmes. L’Anthropologie, 104, 131–145.

    Article  Google Scholar 

  • Debénath, A., Raynal, J. P., Roche, J., Texier, J. P., & Ferembach, D. (1986). Stratigraphie, habitat, typologie et devenir de l’Atérien marocain: Données récentes. L’Anthropologie, 90, 233–246.

    Google Scholar 

  • de Heinzelin, J. (1962). Ishango. Scientific American, 206, 105–116.

    Article  Google Scholar 

  • deMenocal, P. B. (1995). Plio-Pleistocene African climate. Science, 270, 53–59.

    Article  Google Scholar 

  • deMenocal, P. B. (2004). African climate change and faunal evolution during the Pliocene-Pleistocene. Earth and Planetary Science Letters, 220, 3–24.

    Article  Google Scholar 

  • deMenocal, P., Ortiz, J., Guilderson, T., Adkins, J., Sarnthein, M., Baker, L., et al. (2000). Abrupt onset and termination of the African humid period: Rapid climate responses to gradual insolation forcing. Quaternary Science Reviews, 19, 347–361.

    Article  Google Scholar 

  • d’Errico, F., Vanhaeren, M., & Wadley, L. (2008). Possible shell beads from the Middle Stone Age layers of Sibudu Cave, South Africa. Journal of Archaeological Science, 35, 2675–2685.

    Article  Google Scholar 

  • d’Errico, F., Moreno, R. G., & Rifkin, R. F. (2012). Technological, elemental and colorimetric analysis of an engraved ochre fragment from the Middle Stone Age levels of Klasies River Cave 1, South Africa. Journal of Archaeological Science, 39, 942–952.

    Article  Google Scholar 

  • de Ruiter, D. J., Brophy, J. K., Lewis, P. J., Churchill, S. E., & Berger, L. R. (2008). Faunal assemblage composition and paleoenvironment of Plovers Lake, a Middle Stone Age locality in Gauteng Province, South Africa. Journal of Human Evolution, 55, 1102–1117.

    Article  Google Scholar 

  • de Villiers, H. (1973). Human skeletal remains from Border Cave, Ingwavuma District, KwaZulu, South Africa. Annals of the Transvaal Museum, 28, 229–256.

    Google Scholar 

  • de Villiers, H. (1975). A second adult human mandible from Border Cave, Ingwavuma District, KwaZulu, South Africa. South African Journal of Science, 72, 212–215.

    Google Scholar 

  • de Villiers, H., & Fatti, L. P. (1982). The antiquity of the Negro. South African Journal of Science, 72, 212–215.

    Google Scholar 

  • De Vivo, B., Rolandi, G., Gans, P. B., Calvert, A., Bohrson, W. A., Spera, F. J., & Belkin, H. E. (2001). New constraints on the pyroclastic eruptive history of the Campanian volcanic Plain (Italy). Mineralogy and Petrology, 73, 47–65.

    Article  Google Scholar 

  • Domínguez-Rodrigo, M., Mabulla, A., Luque, L., Thompson, J. W., Rink, J., Bushozi, P., et al. (2008). A new archaic Homo sapiens fossil from Lake Eyasi, Tanzania. Journal of Human Evolution, 54, 899–903.

    Article  Google Scholar 

  • Douka, K., Grün, R., Jacobs, Z., Lane, C., Farr, L., Hunt, C., et al. (2014). The chronostratigraphy of the Haua Fteah cave (Cyrenaica, northeast Libya). Journal of Human Evolution, 66, 39–63.

    Article  Google Scholar 

  • Drake, N., & Bristow, C. (2006). Shorelines in the Sahara: Geomorphological evidence for an enhanced monsoon from paleolake Megachad. The Holocene, 16, 901–911.

    Article  Google Scholar 

  • Drennan, M. R. (1929). An Australoid skull from the Cape Flats. Journal of the Royal Anthropological Institute, 59, 417–427.

    Google Scholar 

  • Duarte, C., Mauríco, J., Pettitt, P. B., Souto, P., Trinkaus, E., van der Plicht, H., et al. (1999). The early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern human emergence in Iberia. Proceedings of the National Academy of Sciences of the United States of America, 96, 7604–7609.

    Article  Google Scholar 

  • Dupont, L. (2011). Orbital scale vegetation change in Africa. Quaternary Science Reviews, 30, 3589–3602.

    Google Scholar 

  • Dusseldorp, G. L. (2010). Prey choice during the South African Middle Stone Age: Avoiding dangerous prey or maximizing returns? African Archaeological Review, 27, 107–133.

    Article  Google Scholar 

  • Elhaik, E., Tatarinova, T. V., Klyosov, A. A., & Graur, D. (2014). The ‘extremely ancient’ chromosome that isn’t: A forensic bioinformatics investigation of Albert Perry’s X-degenerate portion of the Y chromosome. European Journal of Human Genetics, 2014, 1–6. doi:10.1038/ejhg.2013.303.

    Google Scholar 

  • Emiliani, C. (1955). Pleistocene temperatures. Journal of Geology, 63, 539–578.

    Article  Google Scholar 

  • Emiliani, C. (1966). Isotopic paleotemperatures. Science, 154, 851–857.

    Article  Google Scholar 

  • Emiliani, C., Cardini, L., Mayeda, T., McBurney, C. B. M., & Tongiorgi, E. (1964). Paleotemperature analysis of fossil shells of marine molluscs (food refuse) from the Arene Candide Cave, Italy, and the Haua Fteah Cave, Cyrenaica. In H. Craig (Ed.), Isotopic and cosmic chemistry (pp. 133–156). Amsterdam: North-Holland.

    Google Scholar 

  • Ennouchi, E. (1962). Un Néanderthalien: L’homme du Jebel Irhoud (Maroc). L’Anthropologie, 66, 279–299.

    Google Scholar 

  • Ennouchi, E. (1963). Les Néanderthaliens du Jebel Irhoud (Maroc). Comptes Rendus des Séances de l’Académie des Sciences (Paris), 256, 2459–2560.

    Google Scholar 

  • Ennouchi, E. (1968). Le deuxième crâne de l’homme de l’Irhoud. Annals de Paléontologie, 55, 117–128.

    Google Scholar 

  • Ennouchi, E. (1969). Présence d’un enfant Néanderthalien au Jebel Irhoud (Maroc). Annals de Paléontologie, 55, 251–265.

    Google Scholar 

  • Epstein, S., Buchsbaum, R., Lowenstam, H. A., & Urey, H. C. (1951). Carbonate-water isotopic temperature scale. Bulletin of the Geological Society of America, 62, 417–426.

    Article  Google Scholar 

  • Epstein, S., Buchsbaum, R., Lowenstam, H. A., & Urey, H. C. (1953). Revised carbonate-water isotopic temperature scale. Bulletin of the Geological Society of America, 64, 1315–1326.

    Article  Google Scholar 

  • Eriksson, A., & Manica, A. (2011). Effect of ancient population structure on the degree of polymorphism shared between modern human populations and ancient hominins. Proceedings of the National Academy of Sciences of the United States of America, 109, 13956–13960.

    Article  Google Scholar 

  • Faith, J. T., Tryon, C., & Peppe, D. J. (2016). Environmental change, ungulate biogeography, and their implications for early human dispersals in equatorial East Africa. In S. C. Jones & B. A. Stewart (Eds.), Africa from MIS 6-2: Population dynamics and paleoenvironments (pp. 233–245). Dordrecht: Springer.

    Google Scholar 

  • Farrera, I., Harrison, S. P., Prentice, I. C., Ramstein, G., Guiot, J., Bartlein, P. J., et al. (1999). Tropical climates at the Last Glacial Maximum: A new synthesis of terrestrial paleoclimate data. I. Vegetation, lake levels and geochemistry. Climate Dynamics, 15, 823–856.

    Article  Google Scholar 

  • Fatti, L. P. (1986). Discriminant analysis in prehistoric physical anthropology. In R. Singer & J. K. Lundy (Eds.), Variation, culture and evolution in African populations (pp. 27–34). Johannesburg: Witwatersrand University Press.

    Google Scholar 

  • Feathers, J. K. (2002). Luminescence dating in less than ideal conditions: Case studies from Klasies River Main Site and Duinefontein, South Africa. Journal of Archaeological Science, 29, 177–194.

    Article  Google Scholar 

  • Feathers, J. K., & Bush, D. A. (2000). Luminescence dating of Middle Stone Age deposits at Die Kelders. Journal of Human Evolution, 38, 91–119.

    Article  Google Scholar 

  • Ferembach, D. (1962). La Nécropole Épipaléolithique de Taforalt (Maroc Oriental). Étude des Squelettes Humains. Casablanca: Edita-Casablanca.

    Google Scholar 

  • Ferembach, D. (1965). Diagrammes craniens sagittaux, et mensurations individuelles des squelettes Ibéromaurusiens de Taforalt (Maroc Oriental). Travaux du centre de recherches anthropologiques, préhistoriques, et etnographiques, Alger.

    Google Scholar 

  • Ferembach, D. (1976a). Les restes humains Atériens de Temara (Campagne 1975). Bulletins et Mémoires de la Société d’anthropologie de Paris, 3, 175–180.

    Article  Google Scholar 

  • Ferembach, D. (1976b). Les restes humains de la grotte de Dar-es-Soltane 2 (Maroc). Campagne 1975. Bulletins et Mémoires de la Société d’Anthropologie de Paris, 3, 183–193.

    Article  Google Scholar 

  • Ferembach, D. (1998). La crâne Atérien de Témara (Maroc, Atlantique). Bulletin d’Archeologie Marocaine, 18, 19–66.

    Google Scholar 

  • Ferembach, D. (1985). On the origin of the Iberomaurusians. A new hypothesis. Journal of Human Evolution, 14, 393–397.

    Article  Google Scholar 

  • Filippi, M. L., & Talbot, M. R. (2005). The paleolimnology of northern Lake Malawi over the last 25 ka based upon the elemental and stable isotopic composition of sedimentary organic matter. Quaternary Science Reviews, 24, 1303–1328.

    Article  Google Scholar 

  • Fitzsimmons, K. E., Hambach, U., Veres, D., & Iovita, R. (2013). The Campanian Ignimbrite eruption: New data on volcanic ash dispersal and its potential impact on human evolution. PLoS ONE, 8(6), e65839. doi:10.1371/journal.pone.0065839.

    Article  Google Scholar 

  • Foerster, V., Junginger, A., Langkamp, O., Gebru, T., Asrat, A., Umer, M., et al. (2012). Climatic change recorded in the sediments of the Chew Bahir basin, southern Ethiopia, during the last 45,000 years. Quaternary International, 274, 25–37.

    Article  Google Scholar 

  • Foley, R., & Lahr, M. M. (1997). Mode 3 technologies and the evolution of modern humans. Cambridge Archaeology Journal, 7, 3–36.

    Article  Google Scholar 

  • Foley, R. A., Maíllo-Fernández, J. M., & Lahr, M. M. (2013). The Middle Stone Age of the Central Sahara: Biogeographical opportunities and technological strategies in later human evolution. Quaternary International, 300, 153–170.

    Article  Google Scholar 

  • Franciscus, R. G. (2003). Internal nasal floor configuration in Homo with special reference to the evolution of Neandertal facial form. Journal of Human Evolution, 44, 701–729.

    Article  Google Scholar 

  • Frayer, D. W., Wolpoff, M. H., Thorne, A. G., Smith, F. H., & Pope, G. G. (1993). Theories of modern human origins: The paleontological test. American Anthropologist, 95, 14–50.

    Article  Google Scholar 

  • Frayer, D. W., Jelínek, J., Oliva, M., & Wolpoff, M. H. (2006). Aurignacian male crania, jaws, and teeth from the Mladeč caves, Moravia, Czech Republic. In M. Teschler-Nicola (Ed.), Early modern humans at the Moravian Gate: The Mladeč caves and their remains (pp. 185–272). Vienna: Springer.

    Chapter  Google Scholar 

  • Frazer, K. A., & The International HapMap Consortium. (2007). A second generation human haplotype map of over 3.1 million SNPs. Nature, 449, 851–861.

    Article  Google Scholar 

  • Fujita, T., Azuma, Y., Fukuyama, R., Hattori, Y., Yoshida, C., Koida, M., et al. (2004). Runx2 induces osteoblast and chondrocyte differentiation and enhances their migration by coupling with PI3K-Akt signaling. Journal of Cell Biology, 166, 85–95.

    Article  Google Scholar 

  • Gabel, C. (1963). Further human remains from the Central African Later Stone Age. South African Archaeological Bulletin, 18, 40–48.

    Article  Google Scholar 

  • Garcin, Y., Junginger, A., Melnick, D., Olago, D. O., Strecker, M. R., & Trauth, M. H. (2009). Late Pleistocene - Holocene rise and collapse of Lake Suguta, northern Kenya Rift. Quaternary Science Reviews, 28, 911–925.

    Article  Google Scholar 

  • Garrigan, D., & Hammer, M. F. (2006). Reconstructing human origins in the genomic era. Nature Reviews Genetics, 7, 669–680.

    Article  Google Scholar 

  • Garrigan, D., Kingan, S. B., Pilkington, M. M., Wilder, J. A., Cox, M. P., Soodyall, H., et al. (2007). Inferring human population sizes, divergence times and rates of gene flow from mitochondrial, X and Y chromosome resequencing data. Genetics, 177, 2195–2207.

    Article  Google Scholar 

  • Garrod, D. A. E. (1970). Pre-Aurignacian and Amudian: A comparative study of the earliest blade industries of the Near East. In K. Gripp, R. Schütrumpf, & H. Schabedissen (Eds.), Frühe Menschheit und Umwelt (pp. 224–229). Köln: Böhlau Verlag.

    Google Scholar 

  • Gasse, F. (2000). Hydrological changes in the African tropics since the Last Glacial Maximum. Quaternary Science Reviews, 19, 189–211.

    Article  Google Scholar 

  • Gasse, F., Chalié, F., Vincens, A., Williams, M. A. J., & Williamson, D. (2008). Climatic patterns in equatorial and southern Africa from 30,000 to 10,000 years ago reconstructed from terrestrial and near-shore proxy data. Quaternary Science Reviews, 27, 2316–2340.

    Article  Google Scholar 

  • Gathogo, P. N., & Brown, F. H. (2006). Stratigraphy of the Koobi Fora Formation (Pliocene and Pleistocene) in the Ileret region of northern Kenya. Journal of African Earth Sciences, 45, 369–390.

    Article  Google Scholar 

  • Gibbard, P. L., & Head, M. J. (2009). IUGS ratification of the Quaternary system/period and the Pleistocene series/epoch with a base at 2.58 Ma. Quaternaire, 20, 411–412.

    Article  Google Scholar 

  • Gliganic, L. A., Jacobs, Z., Roberts, R. G., Domínquez-Rodrigo, M., & Mabulla, A. Z. P. (2012). New ages for Middle and Later Stone Age deposits at Mumba rockshelter, Tanzania: Optically stimulated luminescence dating of quartz and feldspar grains. Journal of Human Evolution, 62, 533–547.

    Article  Google Scholar 

  • Goldberg, P. (2000). Micromorphology and site formation at Die Kelders Cave 1, South Africa. Journal of Human Evolution, 38, 43–90.

    Article  Google Scholar 

  • Gonder, M. K., Mortensen, H. M., Reed, F. A., de Sousa, A., & Tishkoff, S. A. (2007). Whole-mtDNA genome sequence analysis of ancient African lineages. Molecular Biology and Evolution, 24, 757–768.

    Article  Google Scholar 

  • Gokcumen, O., Zhu, Q., Mulder, L. C. F., Iskow, R. C., Austermann, C., Scharer, C. D., et al. (2013). Balancing selection on a regulatory region exhibiting ancient variation that predates human–Neandertal divergence. PLoS Genetics, 9(4), e1003404. doi:10.1371/journal.pgen.1003404.

    Article  Google Scholar 

  • Goodwin, A. J. H. (1929). Report on the stone implements found with the Cape Flats skull. Journal of the Royal Anthropological Institute, 59, 429–438.

    Google Scholar 

  • Görsdorf, J., & Eiwanger, J. (1998). Radiocarbon datings of Late Paleolithic, Epipaleolithic and Neolithic sites in northeastern Morocco. In J. Evin, C. Oberlin, J. P. Daugas & J. F. Salles (Eds.), Actes du 3e Congrès International sur 14C et Archéologie (Mémoires de la Societé Préhistoire Français No. 26) (pp. 365–369). Paris: Societé Préhistoire Français.

    Google Scholar 

  • Gramly, R. M. (1976). Upper Pleistocene archaeological occurrences at site GvJm22, Lukenya Hill, Kenya. Man, 11, 319–344.

    Article  Google Scholar 

  • Gramly, R. M., & Rightmire, G. P. (1973). A fragmentary cranium and dated Later Stone Age assemblages from Lukenya Hill, Kenya. Man, 8, 571–579.

    Article  Google Scholar 

  • Greene, D. L., & Armelagos, G. J. (1972). The Wadi Halfa Mesolithic population (Research Report No. 11). Amherst, MA: University of Massachusetts.

    Google Scholar 

  • Green, R. E., Malaspinas, A. S., Krause, J., Briggs, A., Johnson, P. L. F., Uhler, C., et al. (2008). A complete Neanderthal mitochondrial genome sequence determined by high throughput sequencing. Cell, 134, 416–426.

    Article  Google Scholar 

  • Green, R. E., Krause, J., Briggs, A. W., Maricic, T., Stenzel, U., Kircher, M., et al. (2010). A draft sequence of the Neanderthal genome. Science, 328, 710–722.

    Article  Google Scholar 

  • Grine, F. E. (1998). Additional human fossils from the Middle Stone Age of Die Kelders Cave 1, South Africa: 1995 excavations. South African Journal of Science, 94, 229–235.

    Google Scholar 

  • Grine, F. E. (2000). Middle Stone Age human fossils from Die Kelders Cave 1, Western Cape Province, South Africa. Journal of Human Evolution, 38, 129–145.

    Article  Google Scholar 

  • Grine, F. E. (2012). Observations on Middle Stone Age human teeth from Klasies River Main Site. South Africa Journal of Human Evolution, 63, 750–758.

    Google Scholar 

  • Grine, F. E., & Klein, R. G. (1985). Pleistocene and Holocene human remains from Equus Cave, South Africa. Anthropology, 8, 55–98.

    Google Scholar 

  • Grine, F. E., & Klein, R. G. (1993). Late Pleistocene human remains from the Sea Harvest site, Saldanha Bay, South Africa. South African Journal of Science, 89, 145–152.

    Google Scholar 

  • Grine, F. E., Klein, R. G., & Volman, T. P. (1991). Dating, archaeology and human fossils from the Middle Stone Age Layers of Die Kelders Cave 1, South Africa. Journal of Human Evolution, 21, 363–395.

    Article  Google Scholar 

  • Grine, F. E., Henshilwood, C. S., & Sealy, J. C. (2000). Human remains from Blombos Cave, South Africa: (1997–1998 excavations). Journal of Human Evolution, 38, 755–765.

    Article  Google Scholar 

  • Grine, F. E., & Henshilwood, C. S. (2002). Additional human remains from Blombos Cave, South Africa: (1999–2000 excavations). Journal of Human Evolution, 42, 293–302.

    Article  Google Scholar 

  • Grine, F. E., Pearson, O. M., Klein, R. G., & Rightmire, G. P. (1998). Additional human fossils from Klasies River Mouth, South Africa. Journal of Human Evolution, 35, 95–107.

    Article  Google Scholar 

  • Grine, F. E., Bailey, R. M., Harvati, K., Nathan, R. P., Morris, A. G., Henderson, G. M., et al. (2007). Late Pleistocene human skull from Hofmeyr, South Africa, and modern human origins. Science, 315, 226–229.

    Article  Google Scholar 

  • Grine, F. E., Gunz, P., Betti-Nash, L., Neubauer, S., & Morris, A. G. (2010). Reconstruction of the Late Pleistocene human skull from Hofmeyr, South Africa. Journal of Human Evolution, 59, 1–15.

    Article  Google Scholar 

  • Grine, F. E., Jacobs, R. L., Reed, K. E., & Plavcan, J. M. (2012). The enigmatic molar from Gondolin, South Africa: Implications for Paranthropus paleobiology. Journal of Human Evolution, 63, 597–609.

    Article  Google Scholar 

  • Gronau, I., Hubisz, M. J., Gulko, B., Danko, C. G., & Siepel, A. (2011). Bayesian inference of ancient human demography from individual genome sequences. Nature Genetics, 43, 1031–U151.

    Article  Google Scholar 

  • Grün, R. (2006). Direct dating of human fossils. Yearbook of Physical Anthropology, 49, 2–48.

    Article  Google Scholar 

  • Grün, R., & Beaumont, P. (2001). Border Cave revisited: A revised ESR chronology. Journal of Human Evolution, 40, 467–482.

    Article  Google Scholar 

  • Grün, R., & Stringer, C. B. (1991). Electron spin resonance dating and the evolution of modern humans. Archaeometry, 33, 153–199.

    Article  Google Scholar 

  • Grün, R., Beaumont, P. B., & Stringer, C. B. (1990a). ESR dating evidence for early modern humans at Border Cave in South Africa. Nature, 344, 537–539.

    Article  Google Scholar 

  • Grün, R., Shackleton, N. J., & Deacon, H. J. (1990b). Electron-spin-resonance dating of tooth enamel from Klasies River Mouth. Current Anthropology, 31, 427–432.

    Article  Google Scholar 

  • Grün, R., Beaumont, P., Tobias, P. V., & Eggins, S. (2003). On the age of Border Cave 5 human mandible. Journal of Human Evolution, 45, 155–167.

    Article  Google Scholar 

  • Guérin, G., Murray, A. S., Jain, M., Thomsen, K. J., & Mercier, N. (2013). How confident are we in the chronology of the transition between Howieson’s Poort and Still Bay? Journal of Human Evolution, 64, 314–317.

    Article  Google Scholar 

  • Gunz, P., Bookstein, F. L., Mitteroecker, P., Stadimayr, A., Seidler, H., & Weber, G. (2009). Early modern human diversity suggests subdivided population structure and a complex out-of-Africa scenario. Proceedings of the National Academy of Sciences of the United States of America, 106, 6094–6098.

    Article  Google Scholar 

  • Gunz, P., Neubauer, S., Maureille, S., & Hublin, J. J. (2010). Brain development after birth differs between Neanderthals and modern humans. Current Biology, 20, R921–R922.

    Article  Google Scholar 

  • Hachi, S. (1996). L’Ibéromaurusien, découverte des fouilles d’Afalou (Bédjaîa, Algérie). L’Anthropologie, 100, 55–76.

    Google Scholar 

  • Haile-Selassie, Y., Asfaw, B., & White, T. D. (2004). Hominid cranial remains from upper Pleistocene deposits at Aduma, Middle Awash, Ethiopia. American Journal of Physical Anthropology, 123, 1–10.

    Article  Google Scholar 

  • Halkett, D. J., Hart, T., Yates, R. J., Volman, T. P., Parkington, J. E., Orton, J., et al. (2003). First excavation of intact Middle Stone Age layers at Ysterfontein, Western Cape Province, South Africa: Implications for Middle Stone Age ecology. Journal of Archaeological Science, 30, 955–971.

    Article  Google Scholar 

  • Hammer, M. F., Karafet, T. M., Redd, A. J., Jarjanazi, H., Santachiara-Benerecetti, S., Soodyall, H., et al. (2001). Hierarchical patterns of global human Y-chromosome diversity. Molecular Biology and Evolution, 18, 1189–1203.

    Article  Google Scholar 

  • Hammer, M. F., Woerner, A. E., Mendez, F. L., Watkins, J. C., & Wall, J. D. (2011). Genetic evidence for archaic admixture in Africa. Proceedings of the National Academy of Sciences of the United States of America, 108, 15123–15128.

    Article  Google Scholar 

  • Harvati, K., & Hublin, J. J. (2012). Morphological continuity of the face in the Late Middle and Late Pleistocene hominins from northwestern Africa: A 3D geometric morphometric analysis. In J. J. Hublin & S. P. McPherron (Eds.), Modern origins: A North African perspective (pp. 179–188). Dordrecht: Springer.

    Chapter  Google Scholar 

  • Harvati, K., Frost, S. R., & McNulty, K. P. (2004). Neanderthal taxonomy reconsidered: Implications of 3D primate models of intra- and interspecific differences. Proceedings of the National Academy of Sciences of the United States of America, 101, 1147–1152.

    Google Scholar 

  • Harvati, K., Gunz, P., & Grigorescu, D. (2007). Cioclovina (Romania): Affinities of an early modern European. Journal of Human Evolution, 53, 732–746.

    Article  Google Scholar 

  • Harvati, K., Stringer, C., Grün, R., Aubert, M., Allsworth-Jones, P., & Folorunso, C. A. (2011). The Later Stone Age calvaria from Iwo Eleru, Nigeria: Morphology and chronology. PLoS ONE, 6(9), e24024. doi:10.1371/journal.pone.0024024.

    Article  Google Scholar 

  • Harvati, K., Bauer, C. C., Grine, F. E., Benazzi, S., Ackermann, R. R., van Niekerk, K. L., & Henshilwood, C. S. (2015). A human deciduous molar from the Middle Stone Age (Howiesons Poort) of Klipdrift Shelter, South Africa. Journal of Human Evolution, 82, 190–196.

    Google Scholar 

  • Haughton, S. H. (1917). Preliminary note on the ancient human skull remains from the Transvaal. Transactions of the Royal Society of South Africa, 6, 1–14.

    Article  Google Scholar 

  • Hay, R. L. (1987). Geology of the Laetoli area. In M. D. Leakey & J. M. Harris (Eds.), Laetoli: A Pliocene site in northern Tanzania (pp. 23–47). Oxford: Oxford University Press.

    Google Scholar 

  • Hays, J. D., Imbrie, J., & Shackleton, N. J. (1976). Variation in the earth’s orbit: Pacemaker of the ice ages. Science, 194, 1121–1132.

    Article  Google Scholar 

  • Hearty, P. J., Hollin, J. T., Neumann, A. C., O’Leary, M. J., & McCulloch, M. (2007). Global sea- level fluctuations during the Last Interglaciation (MIS 5e). Quaternary Science Reviews, 26, 2090–2112.

    Article  Google Scholar 

  • Hedges, R. E. M. (2002). Bone diagenssis: An overview of processes. Archaeometry, 44, 319–328.

    Article  Google Scholar 

  • Hendey, Q. B. (1984). Southern African Late Tertiary vertebrates. In R. G. Klein (Ed.), Southern African prehistory and paleoenvironments (pp. 81–106). Rotterdam: A.A. Balkema.

    Google Scholar 

  • Henn, B. M., Gignoux, C. R., Jobin, M., Granka, J. M., Macpherson, J. M., Kidd, J. M., et al. (2011). Hunter-gatherer genomic diversity suggests a southern African origin for modern humans. Proceedings of the National Academy of Sciences of the United States of America, 108, 5154–5162.

    Article  Google Scholar 

  • Henn, B. M., Cavalli-Sforza, L. L., & Feldman, M. W. (2012). The great human expansion. Proceedings of the National Academy of Sciences of the United States of America, 109, 17758–17764.

    Article  Google Scholar 

  • Henshilwood, C. S. (2012). The Still Bay and Howiesons Poort: ‘Paleolithic’ techno-traditions in southern Africa. Journal of World Prehistory, 25, 205–237.

    Article  Google Scholar 

  • Henshilwood, C. S., Sealy, J. C., Yates, R. J., Cruz-Uribe, K., Goldberg, P., Grine, F. E., et al. (2001). Blombos Cave, southern Cape, South Africa: Preliminary report on the 1992–1999 excavations of the Middle Stone Age levels. Journal of Archaeological Science, 28, 421–448.

    Article  Google Scholar 

  • Henshilwood, C. S., d’Errico, F., Yates, R., Jacobs, Z., Tribolo, C., Duller, G. A. T., et al. (2002). Emergence of modern human behaviour: Middle Stone Age engravings from South Africa. Science, 295, 1278–1280.

    Article  Google Scholar 

  • Henshilwood, C. S., d’Errico, F., Vanhaeren, M., van Niekerk, K., & Jacobs, Z. (2004). Middle Stone Age shell beads from South Africa. Science, 384, 404.

    Article  Google Scholar 

  • Henshilwood, C. S., d’ Errico, F., & Watts, I. (2009). Engraved ochres from the Middle Stone Age levels at Blombos Cave, South Africa. Journal of Human Evolution, 57, 27–47.

    Article  Google Scholar 

  • Henshilwood, C. S., d’Errico, F., van Niekerk, K. L., Coquinot, Y., Jacobs, Z., Lauritzen, S. E., et al. (2011). A 100,000-year-old ochre-processing workshop at Blombos Cave, South Africa. Science, 334, 219–222.

    Article  Google Scholar 

  • Henshilwood, C. S., van Niekerk, K. L., Wurz, S., Delagnes, A., Armitage, S. J., Rifkin, R. F., et al. (2014). Klipdrift Shelter, southern Cape, South Africa: Preliminary report on the Howiesons Poort layers. Journal of Archaeological Science, 45, 284–303.

    Article  Google Scholar 

  • Hessler, I., Steinke, S., Groeneveld, J., Dupont, L., & Wefer, G. (2011). Impact of abrupt climate change in the tropical southeast Atlantic during Marine Isotope Stage (MIS) 3. Paleooceanography, 26, PA4209. doi:10.1029/2011PA002118.

  • Hetherington, R., Wiebe, E., Weaver, A. J., Carto, S. L., Eby, M., & MacLoed, R. (2008). Climate, African and Beringian subaerial continental shelves, and migration of early peoples. Quaternary International, 183, 83–101.

    Article  Google Scholar 

  • Hewes, G. W., Irwin, H., Papworth, M., & Saxe, A. (1964). A new fossil human population from the Wadi Halfa area, Sudan. Nature, 203, 341–343.

    Article  Google Scholar 

  • Higham, T. F. G., Jacobi, R. M., & Bronk Ramsey, C. (2006). AMS radiocarbon dating of ancient bone using ultrafiltration. Radiocarbon, 48, 179–195.

    Google Scholar 

  • Hilgren, F. J. (1991). Astronomical calibration of Gauss to Matuyama sapropels in the Mediterranean and implications for the geomagnetic polarity time scale. Earth and Planetary Science Letters, 104, 226–244.

    Article  Google Scholar 

  • Hinz, E. A., & Kohn, M. J. (2010). The effect of tissue structure and soil chemistry on trace element uptake in fossils. Geochimica et Cosmochimica Acta, 74, 3213–3323.

    Article  Google Scholar 

  • Hodgson, J. A., Bergey, C. M., & Disotell, T. R. (2010). Neanderthal genome: The ins and outs of African genetic diversity. Current Biology, 20, R517–R519.

    Article  Google Scholar 

  • Hoefs, J. (2009). Stable isotope geochemistry (6th ed.). Berlin: Springer.

    Google Scholar 

  • Hoelzmann, P., Keding, B., Berke, H., Kröpelin, S., & Kruse, H. J. (2002). Environmental change and archaeology: Lake evolution and human occupation in the eastern Sahara during the Holocene. Paleogeography, Paleoclimatology, Paleoecology, 169, 193–217.

    Article  Google Scholar 

  • Holliday, T. W. (2015). Population affinities of the Jebel Sahaba skeletal sample: Limb proportion evidence. International Journal of Osteoarchaeology, 25, 466–476.

    Google Scholar 

  • Houghton, K., & Thackeray, J. F. (2011). Morphometric comparisons between crania of Late Pleistocene Homo sapiens from Border Cave (BC 1), Tuinplaas (TP 1) and modern southern African populations. Transactions of the Royal Society of South Africa, 66, 159–162.

    Article  Google Scholar 

  • Howell, F. C. (1978). Hominidae. In V. J. Maglio & H. B. S. Cooke (Eds.), Evolution of African mammals (pp. 154–248). Cambridge, MA: Harvard University Press.

    Google Scholar 

  • Hublin, J. J. (1992). Recent human evolution in northwestern Africa. Philosophical Transactions of the Royal Society of London. Series B, Biological sciences, 337, 185–191.

    Article  Google Scholar 

  • Hublin, J. J. (1993). Recent human evolution in northwestern Africa. In M. J. Aitken, C. B. Stringer, & P. Mellars (Eds.), The origin of modern humans and the impact of chronometric dating (pp. 118–131). Princeton: Princeton University Press.

    Google Scholar 

  • Hublin, J. J. (2000). Modern-nonmodern hominid interactions: A Mediterranean perspective. In O. Bar-Yosef & D. R. Pilbeam (Eds.), The geography of Neandertals and modern humans in Europe and the greater Mediterranean (pp. 157–182). Cambridge, MA: Peabody Museum of Archaeology and Ethnology.

    Google Scholar 

  • Hublin, J. J. (2001). Northwestern African Middle Pleistocene hominids and their bearing on the emergence of Homo sapiens. In L. Barham & K. Robson-Brown (Eds.), Human roots. Africa and Asia in the Middle Pleistocene (pp. 99–121). Bristol: Western Academic and Specialist Press.

    Google Scholar 

  • Hublin, J. J., Verna, C., Bailey, S., Smith, T., Olejniczak, A., & Sbihi-Alaoui, F. Z. (2012). Dental evidence from the Aterian human populations of Morocco. In J. J. Hublin & S. P. McPherron (Eds.), Modern origins: A North African perspective (pp. 189–204). Dordrecht: Springer.

    Chapter  Google Scholar 

  • Hughes, A. R. (1990). The Tuinplaas human skeleton from the Springbok Flats, Transvaal. In G. H. Sperber (Ed.), From apes to angels: Essays in honour of Phillip (Vol. Tobias, pp. 197–214). New York: Wiley-Liss.

    Google Scholar 

  • Humphrey, L. T., & Bocaege, E. (2008). Tooth evulsion in the Maghreb: Chronological and geographical patterns. African Archaeological Review, 25, 109–123.

    Article  Google Scholar 

  • Humphrey, L., Bello, S. M., Turner, E., Bouzouggar, A., & Barton, N. (2012). Iberomaurusian funerary behaviour: Evidence from Grotte des Pigeons, Taforalt, Morocco. Journal of Human Evolution, 62, 261–273.

    Article  Google Scholar 

  • Hunt, C., Davison, J., Inglis, R., Farr, L., Reynolds, T., Simpson, D., et al. (2010). Site formation processes in caves: The Holocene sediments of the Haua Fteah, Cyrenaica, Libya. Journal of Archaeological Science, 37, 1600–1611.

    Article  Google Scholar 

  • Imbrie, J. D., & McIntyre, A. (2006). SPECMAP time scale developed by Imbrie et al. 1984 based on normalized planktonic records (normalized O-18 vs time, specmap.017). doi:10.1594/PANGAEA.441706.

  • Imbrie, J., Hays, J. D., Martinson, D. G., McIntyre, A., Mix, A. C., Morley, J. J., et al. (1984). The orbital theory of Pleistocene climate: Support from a revised chronology of the marine δ18O record. In A. Berger (Ed.), Milankovitch and climate, Part 1 (pp. 269–305). Hingham, MA: D. Reidel.

    Google Scholar 

  • Imbrie, J., Berger, A., Boyle, E. A., Clemens, S. C., Duffy, A., Howard, W. R., et al. (1993). On the structure and origin of major glaciations cycles. 2. The 100 000-year cycle. Paleoceanography, 8, 699–735.

    Article  Google Scholar 

  • Indriati, E., Swisher, C. C., Lepre, C., Quinn, R. L., Suriyanto, R. A., Hascaryo, A. T., et al. (2011). The age of the 20 meter Solo River terrace, Java, Indonesia and the survival of Homo erectus in Asia. PLoS ONE, 6(6), e21562. doi:10.1371/journal.pone.0021562.

    Article  Google Scholar 

  • Ingman, M., & Gyllensten, U. (2001). Analysis of the complete human mtDNA genome: Methodology and inferences for human evolution. Journal of Heredity, 92, 454–461.

    Article  Google Scholar 

  • Ingman, M., Kaessmann, H., Pääbo, S., & Gyllensten, U. (2000). Mitochondrial genome variation and the origin of modern humans. Nature, 408, 708–713.

    Article  Google Scholar 

  • Irish, J. (2000). The Iberomaurusian enigma: North Africa progenitor or dead end? Journal of Human Evolution, 39, 393–410.

    Article  Google Scholar 

  • Irish, J. D. (2005). Population continuity vs. discontinuity revisited: Dental affinities among Late Paleolithic through Christian-Era Nubians. American Journal of Physical Anthropology, 128, 520–535.

    Article  Google Scholar 

  • Irish, J. D., & Turner, C. G. (1990). West African dental affinity of Late Pleistocene Nubians: Peopling of the Eurafrican-South Asian triangle, II. Homo, 41, 42–53.

    Google Scholar 

  • Jacobs, Z. (2010). An OSL chronology for the sedimentary deposits from Pinnacle Point Cave 13B—a punctuated presence. Journal of Human Evolution, 59, 289–305.

    Article  Google Scholar 

  • Jacobs, Z., Wintle, A. G., & Duller, G. A. T. (2003a). Optical dating of dune sand from Blombos Cave, South Africa: I—multiple grain data. Journal of Human Evolution, 44, 599–612.

    Article  Google Scholar 

  • Jacobs, Z., Duller, G. A. T., & Wintle, A. G. (2003b). Optical dating of dune sand from Blombos Cave, South Africa: II—single grain data. Journal of Human Evolution, 44, 613–625.

    Article  Google Scholar 

  • Jacobs, Z., Duller, G. A. T., Henshilwood, C. S., & Wintle, A. G. (2006). Extending the chronology of deposits at Blombos Cave, South Africa, back to 140 ka using optical dating of single and multiple grains of quartz. Journal of Human Evolution, 51, 255–273.

    Article  Google Scholar 

  • Jacobs, Z., Roberts, R. G., Galbraith, R. F., Deacon, H. J., Grün, R., Mackay, A., et al. (2008a). Ages for the Middle Stone Age of southern Africa: Implications for human behavior and dispersal. Science, 322, 733–735.

    Article  Google Scholar 

  • Jacobs, Z., Wintle, A. G., Duller, G. A. T., Roberts, R. G., & Wadley, L. (2008b). New ages for the post-Howiesons Poort, late and final Middle Stone Age at Sibudu Cave, South Africa. Journal of Archaeological Science, 35, 1790–1807.

    Article  Google Scholar 

  • Jacobs, Z., Meyer, M. C., Roberts, R. G., Aldeias, V., Dibble, H., & El Hajraoui, M. A. (2011). Single-grain OSL dating at La Grotte des Contrebandiers (‘Smugglers’ Cave’), Morocco: Improved age constraints for the Middle Paleolithic levels. Journal of Archaeological Science, 38, 3631–3643.

    Article  Google Scholar 

  • Jacobs, Z., Hayes, E. H., Roberts, R. G., Galbraith, R. F., & Henshilwood, C. S. (2013). An improved OSL chronology for the Still Bay layers at Blombos Cave, South Africa: Further tests of single-grain dating procedures and a re-evaluation of the timing of the Still Bay industry across southern Africa. Journal of Archaeological Science, 40, 579–594.

    Article  Google Scholar 

  • Jelinek, A. J. (1990). The Amudian in the context of the Mugharan tradition at the Tabun cave (Mt. Carmel), Israel. In P. Mellars (Ed.), The emergence of modern humans (pp. 81–90). Edinburgh: Edinburgh University Press.

    Google Scholar 

  • Jerardino, A., & Marean, C. W. (2010). Shellfish gathering, marine paleoecology and modern human behavior: Perspectives from cave PP13B, Pinnacle Point, South Africa. Journal of Human Evolution, 59, 412–424.

    Article  Google Scholar 

  • Johnson, T. C., Brown, E. T., McManus, J., Barry, S., Barker, P., & Gasse, F. (2002). A high-resolution paleoclimate record spanning the past 25,000 years in southern east Africa. Science, 296, 113–116.

    Article  Google Scholar 

  • Jones, S., Antoniadou, A., Barton, H., Drake, N., Farr, L., Hunt, C., et al. (2016). Patterns of hominin occupation and cultural diversity across the Gebel Akhdar of northern Libya over the last ~ 200 kyr. In S. C. Jones & B. A. Stewart (Eds.), Africa from MIS 6-2: Population dynamics and paleoenvironments (pp. 77–99). Dordrecht: Springer.

    Google Scholar 

  • Jones, T. R. (1940). Human skeletal remains from the Mumbwa Cave, Northern Rhodesia. South African Journal of Science, 37, 313–319.

    Google Scholar 

  • Jouzel, J., Lorius, C., Petit, J. R., Genthon, C., Barkov, N. I., Kotlyakov, V. M., et al. (1987). Vostok Ice Core—a continuous isotope temperature record over the last climatic cycle (160,000 years). Nature, 329, 403–408.

    Article  Google Scholar 

  • Kennett, J. P., Roark, E. B., Cannariato, K. G., Ingram, B. L., & Tada, R. (2000). Latest Quaternary paleoclimatic and radiocarbon chronology, Hole 1017E, southern California margin. In M. Lyle, I. Koizumi, C. Richter & T. C. Moore (Eds.), Proceedings of the ocean drilling program, scientific results (Vol. 167) (pp. 249–254). College Station, TX: Ocean Drilling Program.

    Google Scholar 

  • Kidd, J. M., Gravel, S., Byrnes, J., Moreno-Estrada, A., Musharoff, S., Bryc, K., et al. (2012). Population genetic inference from personal genome data: Impact of ancestry and admixture on human genomic variation. American Journal of Human Genetics, 91, 660–671.

    Article  Google Scholar 

  • Kivisild, T., Shen, P., Wall, D. P., Do, B., Sung, R., Davis, K., et al. (2006). The role of selection in the evolution of human mitochondrial genomes. Genetics, 172, 373–387.

    Article  Google Scholar 

  • Klein, R. G. (1970). Problems in the study of the Middle Stone Age of South Africa. South African Archaeological Bulletin, 25, 127–135.

    Article  Google Scholar 

  • Klein, R. G. (1976). The mammalian fauna of the Klasies River Mouth sites, southern Cape Province, South Africa. South African Archaeology Bulletin, 21, 75–98.

    Article  Google Scholar 

  • Klein, R. G. (1983). The Stone Age prehistory of southern Africa. Annual Review of Anthropology, 12, 25–48.

    Article  Google Scholar 

  • Klein, R. G. (1984). Later Stone Age faunal samples from Heuningneskrans Shelter (Transvaal) and Leopard’s Hill Cave (Zambia). South African Archaeological Bulletin, 39, 109–116.

    Article  Google Scholar 

  • Klein, R. G. (1989). Why does skeletal part representation differ between smaller and larger bovids at Klasies River Mouth and other archeological sites? Journal of Archaeological Science, 16, 363–381.

    Article  Google Scholar 

  • Klein, R. G. (2009). The human career: Human biological and cultural origins (3rd ed.). Chicago: The University of Chicago Press.

    Book  Google Scholar 

  • Klein, R. G., & Scott, K. (1986). Re-analysis of the faunal assemblages from Haua Fteah and other Late Quaternary archaeological sites in Cyrenaican Libya. Journal of Archaeological Science, 13, 515–542.

    Article  Google Scholar 

  • Klein, R. G., & Cruz-Uribe, K. (1996). Exploitation of large bovids and seals and Middle and Later Stone Age sites in South Africa. Journal of Human Evolution, 31, 315–334.

    Article  Google Scholar 

  • Klein, R. G., Cruz-Uribe, K., & Beaumont, P. B. (1991). Environmental, ecological, and paleoanthropological implications of the Late Pleistocene mammalian fauna from Equus Cave, Northern Cape Province, South Africa. Quaternary Research, 36, 94–119.

    Article  Google Scholar 

  • Klein, R. G., Avery, G., Cruz-Uribe, K., Halkett, D. J., Parkington, J. E., Steele, T., et al. (2004). The Ysterfontein 1 Middle Stone Age site, South Africa, and early human exploitation of coastal resources. Proceedings of the National Academy of Sciences of the United States of America, 101, 5708–5715.

    Article  Google Scholar 

  • Komori, T. (2010). Regulation of osteoblast differentiation by Runx2. Advances in Experimental Medicine and Biology, 658, 43–49.

    Article  Google Scholar 

  • Krause, J., Fu, Q., Good, J. M., Viola, B., Shunkov, M. V., Derevianko, A. P., & Pääbo, S. (2010). The complete mitochondrial DNA genome of an unknown hominin from southern Siberia. Nature, 464, 894–897.

    Article  Google Scholar 

  • Kröpelin, S., Verschuren, D., Lézine, A. M., Eggermont, H., Cocquyt, C., Francus, P., et al. (2008). Climate-driven ecosystem succession in the Sahara: The past 6,000 years. Science, 320, 765–768.

    Article  Google Scholar 

  • Kuman, K., Inbar, M., & Clarke, R. J. (1999). Paleoenvironments and cultural sequence of the Florisbad Middle Stone Age hominid site, South Africa. Journal of Archaeological Science, 26, 1409–1425.

    Article  Google Scholar 

  • Kuper, R., & Kröpelin, S. (2006). Climate-controlled Holocene occupation of the Sahara: Motor of Africa’s evolution. Science, 313, 803–807.

    Article  Google Scholar 

  • Kusimba, S. B. (2001). The early later Stone Age in East Africa: Excavations and lithic assemblages from Lukenya Hill. African Archaeological Review, 18, 77–123.

    Article  Google Scholar 

  • Lachance, J., Vernot, B., Elbers, C. C., Ferwerda, B., Froment, A., Bodo, J. M., et al. (2012). Evolutionary history and adaptation from high-coverage whole-genome sequences of diverse African hunter-gatherers. Cell, 150, 457–469.

    Article  Google Scholar 

  • Lahr, M. M. (1996). The evolution of modern human diversity. A study of cranial variation. Cambridge: Cambridge University Press.

    Google Scholar 

  • Lalueza-Fox, C., & Gilbert, M. T. P. (2011). Paleogenomics of archaic hominins. Current Biology, 21, R1002–R1009. doi:10.1016/j.cub.2011.11.021.

    Article  Google Scholar 

  • Lam, Y. M., Pearson, O. M., & Smith, C. M. (1996). Chin morphology and sexual dimorphism in the fossil hominid mandible sample from Klasies River Mouth. American Journal of Physical Anthropology, 100, 545–557.

    Article  Google Scholar 

  • Landais, A. J., Barnola, M., Masson-Delmotte, V., Jouzel, J., Chappellaz, J., Caillon, N., et al. (2004). A continuous record of temperature evolution over a sequence of Dansgaard-Oeschger events during Marine Isotopic Stage 4 (76 to 62 kyr BP). Geophysical Research Letters, 31, L22211. doi:10.1029/2004GL021193.

    Article  Google Scholar 

  • Lane, C., Barker, G., Reynolds, T., Inglis, R., Barton, N., Albert, P., et al. (2011). Dating the Paleolithic occupation of the Haua Fteah cave, Libya, using cryptotephrochronology (Abstract 1889). XVIII INQUA Congress, Quaternary Sciences—the view from the mountains. Bern, Switzerland.

    Google Scholar 

  • Lane, C. S., Chorn, B. T., & Johnson, T. C. (2013). Ash from the Toba supereruption in Lake Malawi shows no volcanic winter in East Africa at 75 ka. Proceedings of the National Academy of Sciences of the United States of America, 110, 8025–8029.

    Article  Google Scholar 

  • Lang, N., & Wolff, E. W. (2011). Interglacial and glacial variability from the last 800 ka in marine, ice and terrestrial archives. Climate of the Past, 7, 361–380.

    Article  Google Scholar 

  • Larrasoaña, J. C. (2012). A northeast Saharan perspective on environmental variability in North Africa and its implications for modern human origins. In J. J. Hublin & S. P. McPherron (Eds.), Modern origins: A North African perspective (pp. 19–34). Dordrecht: Springer.

    Chapter  Google Scholar 

  • Leakey, R. E. F. (1969). Early Homo sapiens remains from the Omo River region of south-west Ethiopia. Nature, 222, 1132–1133.

    Article  Google Scholar 

  • Lebamba, J., Vincens, A., & Maley, J. (2012). Pollen, vegetation change and climate at Lake Barombi Mbo (Cameroon) during the last c. 33 000 cal yr BP: A numerical approach. Climate of the Past, 8, 59–78.

    Article  Google Scholar 

  • Li, H., & Durbin, R. (2011). Inference of human population history from individual whole-genome sequences. Nature, 475, 403–407.

    Article  Google Scholar 

  • Li, J. Z., Absher, D. M., Tang, H., Southwick, A. M., Casto, A. M., Ramachandran, S., et al. (2008). Worldwide human relationships inferred from genome-wide patterns of variation. Science, 319, 1100–1104.

    Article  Google Scholar 

  • Linz, B., Balloux, F., Moodley, Y., Manica, A., Liu, H., Roumagnac, P., et al. (2007). An African origin for the intimate association between humans and Helicobacter pylori. Nature, 445, 915–918.

    Article  Google Scholar 

  • Lisiecki, L. E., & Raymo, M. E. (2005). A Plio-Pleistocene stack of 57 globally distributed benthic δ18O records. Paleoceanography, 20, PA2007. doi:10.1029/2005PA001164.

    Google Scholar 

  • Lombard, M., & Phillipson, L. (2010). Indications of bow and stone-tipped arrow use 64,000 years ago in KwaZulu-Natal, South Africa. Antiquity, 84, 635–648.

    Article  Google Scholar 

  • Louw, A. W., Brain, C. K., Vogel, J. C., Mason, R. J., & Eloff, J. F. (1969). Bushman Rock Shelter, Origstad, Eastern Transvaal: A preliminary investigation. South African Archaeological Bulletin, 24, 39–60.

    Article  Google Scholar 

  • Lowe, J., Barton, N., Blockley, S., Bronk Ramsey, C., Cullen, V. L., Davies, W., et al. (2012). Volcanic ash layers illuminate the resilience of Neanderthals and early modern humans to natural hazards. Proceedings of the National Academy of Sciences of the United States of America, 109, 13532–13537.

    Article  Google Scholar 

  • Lubell, D. (1974). The Fakhurian. A Late Paleolithic industry from Upper Egypt (Geological Survey of Egypt, Paper No. 58). Cairo: The Geological Survey of Egypt.

    Google Scholar 

  • Lubell, D. (1984). Paleoenvironments and Epi-paleolithic economies in the Maghreb (20000 to 5000 B.P.). In J. D. Clark & S. A. Brandt (Eds.), From hunters to farmers: The causes and consequences of food production in Africa. (pp. 41–56). Berkeley: University of California Press.

    Google Scholar 

  • Lubell, D. (2001). Late Pleistocene—Early Holocene Maghreb. In P. N. Peregrine & M. Ember (Eds.), Encyclopedia of prehistory (Vol. 1, pp. 129–149)., Africa New York: Kluwer.

    Google Scholar 

  • Lyons, R. P., Scholz, C. A., Buoniconti, M. R., & Martin, M. R. (2011). Late Quaternary stratigraphic analysis of the Lake Malawi Rift, East Africa: An integration of drill-core and seismic-reflection data. Paleogeography, Paleoclimatology, Paleoecology, 303, 20–37.

    Article  Google Scholar 

  • Magori, C. C., & Day, M. H. (1983). Laetoli Hominid 18: An early Homo sapiens skull. Journal of Human Evolution, 12, 747–754.

    Article  Google Scholar 

  • Manega, P. C. (1993). Geochronology, geochemistry, and isotopic study of the Plio-Pleistocene hominid sites and the Ngorongoro Volcanic Highlands in northern Tanzania. Ph.D. Dissertation, University of Colorado.

    Google Scholar 

  • Manega, P. C. (1995). New geochronological results from the Ndutu, Naisiusu, and Ngaloba Beds at Olduvai and Laetoli in northern Tanzania: Their significance for the evolution of modern humans (Abstract). Preservation and use of Olduvai Gorge, Laetoli, rock art, and other paleoanthropological resources in Tanzania, held at the Bellagio Center, Italy, June 5–9, 1995.

    Google Scholar 

  • Manica, A., Amos, B., Balloux, F., & Hanihara, T. (2007). The effect of ancient population bottlenecks on human phenotypic variation. Nature, 448, 346–348.

    Article  Google Scholar 

  • Marean, C. W. (1992). Implications of late Quaternary mammalian fauna from Lukenya Hill (south-central Kenya) for paleoenvironmental change and faunal extinctions. Quaternary Research, 37, 239–255.

    Article  Google Scholar 

  • Marean, C. W. (2010). Pinnacle Point Cave 13B (Western Cape Province, South Africa) in context: The Cape Floral kingdom, shellfish, and modern human origins. Journal of Human Evolution, 59, 425–443.

    Article  Google Scholar 

  • Marean, C. W., Nilssen, P. J., Brown, K., Jerardino, A., & Stynder, D. (2004). Paleoanthropological investigations of Middle Stone Age sites at Pinnacle Point, Mossel Bay (South Africa): Archaeology and hominid remains from the 2000 field season. Paleoanthropology, 2004, 14–83.

    Google Scholar 

  • Marean, C. W., Bar-Matthews, M., Bernatchez, J., Fisher, E., Goldberg, P., Herries, A. I., et al. (2007). Early human use of marine resources and pigment in South Africa during the Middle Pleistocene. Nature, 449, 905–908.

    Article  Google Scholar 

  • Mariotti, V., Bonfighioli, B., Facchini, F., Condemi, S., & Belcastro, M. G. (2009). Funerary practices of the Iberomaurusian population of Taforalt (Tafoughalt; Morocco, 11-12,000 BP): New hypotheses based on a grave by grave skeletal inventory and evidence of deliberate human modification of the remains. Journal of Human Evolution, 56, 340–354.

    Article  Google Scholar 

  • Mark, D. F., Petraglia, M., Smith, V. C., Morgan, L. E., Barfold, D. N., Ellis, B. S., et al. (2014). A high-precision 40Ar/39Ar age for the Young Toba Tuff and dating of ultra-distal tephra: Forcing of Quaternary climate and implications for hominin occupation of India. Quaternary Geochronology, 21, 90–103.

    Article  Google Scholar 

  • Martinson, D. G., Pisias, N. G., Hays, J. D., Imbrie, J., Moore, Theodore C., et al. (1987). Age dating and the orbital theory of the ice ages: Development of a high resolution 0 to 300,000-year chronostratigraphy. Quaternary Research, 27, 1–29.

    Article  Google Scholar 

  • Mason, R. J. (1969). Tentative interpretations of new radiocarbon dates for stone artefact assemblages from Rose Cottage Cave, O.F.S. and Bushman Rock Shelter, Tvl. South African Archaeological Bulletin, 24, 57–59.

    Article  Google Scholar 

  • McBrearty, S., & Brooks, A. S. (2000). The revolution that wasn’t: A new interpretation of the origin of modern human behavior. Journal of Human Evolution, 39, 453–563.

    Article  Google Scholar 

  • McBurney, C. B. M. (1958). Evidence for the distribution in space and time of Neanderthaloids and allied strains in northern Africa. In G. H. R. von Koenigswald (Ed.), Hundert Jahre Neanderthaler (Neanderthal Centenary 1856-1956) (pp. 253–264). Köln: Böhlau.

    Google Scholar 

  • McBurney, C. B. M. (1967). The Haua Fteah (Cyrenaica) and the Stone Age of the South-East Mediterranean. Cambridge: Cambridge University Press.

    Google Scholar 

  • McBurney, C. B. M., Trevor, J. C., & Wells, L. H. (1953a). A fossil human mandible from a Levalloiso-Mousterian horizon in Cyrenaica. Nature, 172, 889–892.

    Article  Google Scholar 

  • McBurney, C. B. M., Trevor, J. C., & Wells, L. H. (1953b). The Haua Fteah fossil jaw. Journal of the Royal Anthropological Institute, 83, 71–85.

    Google Scholar 

  • McCall, G. S. (2006). Multivariate perspectives on change and continuity in the Middle Stone Age lithics from Klasies River Mouth, South Africa. Journal of Human Evolution, 51, 429–439.

    Article  Google Scholar 

  • McCrea, J. M. (1950). On the isotopic chemistry of carbonates and a paleotemperature scale. Journal of Chemical Physics, 18, 849–857.

    Article  Google Scholar 

  • McCrossin, M. L. (1992). Human molars from later Pleistocene deposits of Witkrans Cave, Gaap escarpment, Kalahari margin. Human Evolution, 7, 1–10.

    Article  Google Scholar 

  • McDermott, F., Stringer, C., Grün, R., Williams, C. T., Din, V. K., & Hawkesworth, C. J. (1996). New late Pleistocene uranium-thorium and ESR dates for the Singa hominid (Sudan). Journal of Human Evolution, 31, 507–516.

    Article  Google Scholar 

  • McDougall, I., & Brown, F. H. (2006). Precise 40Ar/39Ar geochronology for the upper Koobi Fora Formation, Turkana Basin, northern Kenya. Journal of the Geological Society of London, 163, 205–220.

    Article  Google Scholar 

  • McDougall, I., Brown, F. H., & Fleagle, J. G. (2005). Stratigraphic placement and age of modern humans from Kibish, Ethiopia. Nature, 433, 733–736.

    Article  Google Scholar 

  • McDougall, I., Brown, F. H., & Fleagle, J. G. (2008). Sapropels and the age of hominins Omo I and II, Kibish, Ethiopia. Journal of Human Evolution, 55, 409–420.

    Article  Google Scholar 

  • Medig, M., Meier, R., Sahnouni, M., & Derradji, A. (1996). Découverte d’un crâne humain dans les niveaux Ibéromaurusiens de la grotte de Taza I, Jijel, Algérie. Comptes Rendus de l’Academie des Sciences Paris, Série II, Sciences de la Terre et des Planetes, 323, 825–831.

    Google Scholar 

  • Mehlman, M. J. (1984). Archaic Homo sapiens at Lake Eyasi, Tanzania: Recent misrepresentations. Journal of Human Evolution, 13, 487–501.

    Article  Google Scholar 

  • Mehlman, M. J. (1987). Provenience, age and associations of archaic Homo sapiens crania from Lake Eyasi, Tanzania. Journal of Archaeological Science, 14, 133–162.

    Article  Google Scholar 

  • Meier, R. J., Sahnouni, M., Medig, M., & Abdelkader Derradji, A. (2003). Human skull from the Taza locality, Jijel, Algeria. Anthropologischer Anzeiger, 61, 129–140.

    Google Scholar 

  • Meignen, L. (2000). Early Middle Paleolithic blade technology in southwestern Asia. Acta Anthropologica Sinica, 19 (suppl.), 158–168.

    Google Scholar 

  • Meignen, L. (2007). Middle Paleolithic blady assemblages in the Near East: A reassessment. In V. P. Lyubin (Ed.), Caucasus and the initial human dispersals in the Old World (pp. 133–148). Saint-Petersburg: Russian Academy of Sciences.

    Google Scholar 

  • Mendez, F. L., Krahn, T., Schrack, B., Krahn, A. M., Veeramah, K. R., Woerner, A. E., et al. (2013). An African American paternal lineage adds an extremely ancient root to the human Y chromosome phylogenetic tree. American Journal of Human Genetics, 92, 454–459.

    Article  Google Scholar 

  • Mercader, J., & Brooks, A. S. (2001). Across forests and savannas: Later Stone Age assemblages from Ituri and Semliki, Democratic Republic of Congo. Journal of Anthropological Research, 57, 197–217.

    Google Scholar 

  • Mercier, N., & Valladas, H. (2003). Reassessment of TL age estimates of burnt flints from the Paleolithic site of Tabun Cave, Israel. Journal of Human Evolution, 45, 401–409.

    Article  Google Scholar 

  • Meyer, M., Kircher, M., Gansauge, M. T., Li, H., Racimo, F., Mallick, S., et al. (2012). A high-coverage genome sequence from an archaic Denisovan individual. Science, 338, 222–226.

    Article  Google Scholar 

  • Michels, J. W., & Marean, C. W. (1984). A Middle Stone Age occupation site at Porc-Epic Cave, Dire Dawa (east-central Ethiopia), Part II. African Archaeological Review, 2, 64–71.

    Google Scholar 

  • Mikdad, A., & Eiwanger, J. (2000). Recherches préhistoriques et protohistoriques dans le Rif oriental (Maroc). Rapport Préliminaire. Beiträge zur Allgemeinen und Vergleichenden Archäologie, 20, 109–167.

    Google Scholar 

  • Millard, A. R. (2008). A critique of the chronometric evidence for hominid fossils: I. Africa and the Near East 500-50 ka. Journal of Human Evolution, 54, 848–874.

    Article  Google Scholar 

  • Millard, A. R., & Hedges, R. E. M. (1995). The role of the environment in uranium uptake by buried bone. Journal of Archaeological Science, 22, 239–250.

    Article  Google Scholar 

  • Miller, G. H., Beaumont, P. B., Deacon, H. J., Brooks, A. S., Hare, P. E., & Jull, A. J. T. (1999). Earliest modern humans in southern Africa dated by isoleucine epimerization in ostrich eggshell. Quaternary Science Reviews, 18, 1537–1548.

    Article  Google Scholar 

  • Miller, S. F. (1971). The age of the Nachikufan industries in Zambia. South African Archaeological Bulletin, 26, 143–146.

    Article  Google Scholar 

  • Mills, S. C., Grab, S. W., Rea, B. R., Carr, S. J., & Farrow, A. (2012). Shifting westerlies and precipitation patterns during the Late Pleistocene in southern Africa determined using glacier reconstruction and mass balance modelling. Quaternary Science Reviews, 55, 145–159.

    Google Scholar 

  • Milo, R. G. (1998). Evidence for hominid predation at Klasies River Mouth, South Africa, and its implications for the behaviour of early modern humans. Journal of Archaeological Science, 25, 99–133.

    Article  Google Scholar 

  • Minugh-Purvis, N. (1993). Reexamination of the immature hominid maxilla from Tangier, Morocco. American Journal of Physical Anthropology, 92, 449–461.

    Article  Google Scholar 

  • Mitchell, P. (2008). Developing the archaeology of Marine Isotope Stage 3. South African Archaeological Society Goodwin Series, 10, 52–65.

    Google Scholar 

  • Moodley, Y., Linz, B., Bond, R. P., Nieuwoudt, M., Soodyall, H., Schlebusch, C. M., et al. (2012). Age of the association between Helicobacter pylori and man. PLoS Pathogens, 8(5), e1002693.

    Article  Google Scholar 

  • Morgan, L. E., Renne, P. R., Taylor, R. E., & WoldeGabriel, G. (2009). Archaeological age constraints from extrusion ages of obsidian: Examples from the Middle Awash, Ethiopia. Quaternary Geochronology, 4, 193–203.

    Article  Google Scholar 

  • Morris, A. G. (1992). Biological relationships between Upper Pleistocene and Holocene populations in southern Africa. In G. Bräuer & F. H. Smith (Eds.), Continuity or replacement: Controversies in Homo sapiens evolution (pp. 131–143). Rotterdam: Balkema.

    Google Scholar 

  • Morris, A. G. (2002). Isolation and the origin of the Khoisan: Late Pleistocene and early Holocene human evolution at the southern end of Africa. Human Evolution, 17, 105–114.

    Article  Google Scholar 

  • Morris, A. G. (2003). The myth of the East African “Bushmen”. South African Archaeological Bulletin, 58, 85–90.

    Article  Google Scholar 

  • Mourre, V., Villa, P., & Henshilwood, C. (2010). Early use of pressure flaking on lithic artifacts at Blombos Cave, South Africa. Science, 330, 659–662.

    Article  Google Scholar 

  • Nami, M. (2007). Les techno-complexes Ibéromaurusiens d´Ifri El Baroud (Rif Oriental, Maroc). Forschungen zur Archäologie Außereuropäischer Kulturen, 2, 183–239.

    Google Scholar 

  • Nara, M. T. (1994). Etude de la variabilité de certains caractères métriques et morphologiques des Néandertaliens. Ph.D. Dissertation, University of Bordeaux.

    Google Scholar 

  • Noonan, J. P., Coop, G., Kudaravalli, S., Smith, D., Krause, J., Alessi, J., et al. (2006). Sequencing and analysis of Neanderthal genomic DNA. Science, 314, 1113–1118.

    Google Scholar 

  • Orban, R., Semal, P., & Twiesselmann, F. (2001). Sur la biométrie des mandibules et des dents humaines d’Ishango (LSA, République démocratique du Congo). Bulletins et mémoires de la Société d’Anthropologie de Paris, 13(1–2), 1–13.

    Google Scholar 

  • Parkington, J. (1990). A critique of the consensus view on the age of Howieson’s Poort assemblages in South Africa. In P. Mellars (Ed.), The emergence of modern humans (pp. 34–55). Edinburgh: Edinburgh University Press.

    Google Scholar 

  • Parkington, J., & Poggenpoel, C. (1987). Diepkloof Rock Shelter. In J. Parkington & M. Hall (Eds.), Papers in prehistory of the Western Cape, South Africa (BAR International Series No. 332) (pp. 269–293). Oxford: Archaeopress.

    Google Scholar 

  • Partridge, T. C., deMenocal, P. B., Lorentz, S. A., Paiker, M. J., & Vogel, J. C. (1997). Orbital forcing of climate over South Africa: A 200,000-year rainfall record from the Pretoria saltpan. Quaternary Science Reviews, 16, 1125–1133.

    Article  Google Scholar 

  • Partridge, T. C., Scott, L., & Hamilton, J. E. (1999). Synthetic reconstructions of southern African environments during the Last Glacial Maximum (21-18 kyr) and the Holocene Altithermal (8-6 kyr). Quaternary International, 57(58), 207–214.

    Article  Google Scholar 

  • Pearson, O. (2000). Human remains from Twin Rivers. Appendix 8. In L. Barham (Ed.), The Middle Stone Age of Zambia (pp. 281–282). Bristol: Western Academic and Specialist Press.

    Google Scholar 

  • Pearson, O. M. (2011). Integration of the genetic, anatomical and archaeological data for the African origin of modern humans: Problems and prospects. In S. C. Reynolds & A. Gallagher (Eds.), African genesis: Perspectives on hominin evolution (pp. 423–448). Cambridge: Cambridge University Press.

    Google Scholar 

  • Pearson, O. M., & Grine, F. E. (1996). Morphology of the Border Cave hominid ulna and humerus. South African Journal of Science, 92, 231–236.

    Google Scholar 

  • Pearson, O. M., & Grine, F. E. (1997). Re-analysis of the hominid radii from Cave of Hearths and Klasies River Mouth, South Africa. Journal of Human Evolution, 32, 577–592.

    Google Scholar 

  • Pearson, O. M., Churchill, S. E., Grine, F. E., Trinkaus, E., & Holliday, T. W. (1998). Multivariate analyses of the hominid ulna from Klasies River Mouth. Journal of Human Evolution, 34, 653–656.

    Article  Google Scholar 

  • Pearson, O., Grine, F. E., Barham, L., & Stringer, C. (2000). Human remains from the Middle and Later Stone Age of Mumbwa Caves. In L. Barham (Ed.), The Middle Stone Age of Zambia, South Central Africa (pp. 149–164). Bristol, UK: Western Academic & Specialist Press.

    Google Scholar 

  • Pearson, O. M., Royer, D. F., Grine, F. E., & Fleagle, J. G. (2008a). A description of the Omo I postcranial skeleton, including newly discovered fossils. Journal of Human Evolution, 55, 421–437.

    Article  Google Scholar 

  • Pearson, O. M., Fleagle, J. G., Grine, F. E., & Royer, D. F. (2008b). Further new hominin fossils from the Kibish Formation, southwestern Ethiopia. Journal of Human Evolution, 55, 444–447.

    Article  Google Scholar 

  • Peers, B., & Goodwin, A. J. H. (1953). Two caves at Kalk Bay, Cape Peninsula. Part II: Peer’s Shelter B/102. South African Archaeology Bulletin, 8, 67–77.

    Google Scholar 

  • Petit, J. R., Jouzel, J., Raynaud, D., Barkov, N. I., Barnola, J. M., Basile, I., et al. (1999). Climate and atmospheric history of the past 420,000 years from the Vostok ice core, Antarctica. Nature, 399, 429–436.

    Article  Google Scholar 

  • Pfeiffer, S., & Zehr, M. K. (1996). A morophological and histological study of the human humerus from Border Cave. Journal of Human Evolution, 31, 49–59.

    Article  Google Scholar 

  • Pfeiffer, S., Lazenby, R., & Thackeray, J. F. (1996). Tuinplaas: Affinity assessed from long bone cross-sectional geometry. Kaupia, 6, 137–143.

    Google Scholar 

  • Pickford, M. (1986). Cainozoic paleontological sites of Western Kenya. Münchner Geowissenschaftlich Abhandlungen, 8, 1–151.

    Google Scholar 

  • Pickrell, J. K., Patterson, N., Barbieri, C., Berthold, F., Gerlach, L., Güldemann, T., et al. (2012). The genetic prehistory of southern Africa. Nature Communications, 3, 1143. doi:10.1038/ncomms2140.

    Article  Google Scholar 

  • Pike, A. W. G., Eggins, S., Grün, R., & Thackeray, F. (2004). U-series dating of TP1, an almost complete human skeleton from Tuinplaas (Springbok Flats), South Africa. South African Journal of Science, 100, 381–383.

    Google Scholar 

  • Pike, A. W. G., Hedges, R. E. M., & Van Calsteren, P. (2002). U-series dating of bone using the diffusion-adsorption model. Geochimica et Cosmochimica Acta, 66, 4268–4273.

    Article  Google Scholar 

  • Pinhasi, R. (2002). Biometric study of the affinities of NK—a quantitative analysis of mandible dimensions. In P. M. Vermeersch (Ed.), Paleolithic quarrying sites in upper and middle Egypt (pp. 283–336). Leuven: Leuven University Press.

    Google Scholar 

  • Pinhasi, R., & Semal, P. (2000). The position of the Nazlet Khater specimen among prehistoric and modern African and Levantine populations. Journal of Human Evolution, 39, 269–288.

    Article  Google Scholar 

  • Pleurdeau, D. (2005). Human technical behavior in the African Middle Stone Age: The lithic assemblage of Porc-Épic Cave (Dire Dawa, Ethiopia). African Archaeological Review, 22, 177–197.

    Article  Google Scholar 

  • Plug, I. (2004). Resource exploitation: Animal use during the Middle Stone Age at Sibudu Cave, KwaZulu-Natal. South African Journal of Science, 100, 151–158.

    Google Scholar 

  • Pokras, E. M., & Mix, A. C. (1985). Eolian evidence for spatial variability of Late Quaternary climates in tropical Africa. Quaternary Research, 24, 137–149.

    Article  Google Scholar 

  • Ponce de León, M. S., & Zollikofer, C. P. E. (2001). Neanderthal cranial ontogeny and its implications for late hominid diversity. Nature, 412, 534–538.

    Article  Google Scholar 

  • Porter, S. C. (1989). Some geological implications of average Quaternary glacial conditions. Quaternary Research, 32, 245–261.

    Article  Google Scholar 

  • Poznik, G. D., Henn, B. M., Yee, M. C., Sliwerska, E., Euskirchen, G. M., Lin, A. A., et al. (2013). Sequencing Y chromosomes resolves discrepancy in time to common ancestor of males versus females. Science, 341, 562–565.

    Article  Google Scholar 

  • Pritchard, J. K., Seielstad, M. T., Perez-Lezaun, A., & Feldman, M. W. (1999). Population growth of human Y chromosome microsatellites. Molecular Biology and Evolution, 16, 1791–1798.

    Article  Google Scholar 

  • Protsch, R. (1974). The Fish Hoek hominid: Another member of basic Homo sapiens afer. Anthropologischer Anzeiger, 34, 241–249.

    Google Scholar 

  • Protsch, R. (1975). The absolute dating of Upper Pleistocene sub-Saharan fossil hominids and their place in human evolution. Journal of Human Evolution, 4, 297–322.

    Article  Google Scholar 

  • Protsch, R. R. R. (1977). Mumbwa: Its absolute chronology and archaeology. Zeitschrift fur Morphologie und Anthropologie, 68, 1–7.

    Google Scholar 

  • Protsch, R. (1981). The paleoanthropological finds of the Pliocene and Pleistocene. (Die Archaologischen und anthropologischen Ergebnisse der Kohl-Larsen-Expeditionen in Nord-Tanzania 1933-1939). Tübingen: Verlag Archaeologica Venatoria.

    Google Scholar 

  • Protsch, R., & de Villiers, H. (1974). Bushman Rock Shelter, Origstad, Eastern Transvaal, South Africa. Journal of Human Evolution, 3, 387–396.

    Article  Google Scholar 

  • Pycraft, W. P. (1925). On the calvaria found at Boskop, Transvaal, 1913, and its relationship to Cromagnard and negroid skulls. Journal of the Royal Anthropological Institute, 55, 179–198.

    Google Scholar 

  • Rak, Y. (1998). Does any Mousterian cave present evidence of two hominid species? In T. Akazawa, K. Aoki, & O. Bar-Yosef (Eds.), Neandertals and modern humans in western Asia (pp. 353–366). New York: Plenum.

    Google Scholar 

  • Rampino, M. R., & Self, S. (1992). Volcanic winter and accelerated glaciations following the Toba super-eruption. Nature, 359, 50–52.

    Article  Google Scholar 

  • Rampino, M. R., & Self, S. (1993). Climate-volcanism feedback and the Toba eruption of ~74,000 years ago. Quaternary Research, 40, 269–280.

    Article  Google Scholar 

  • Reed, C. A. (1965). A human frontal bone from the Late Pleistocene of the Kom Ombo Plain, Upper Egypt. Man, 65, 101–104.

    Article  Google Scholar 

  • Reich, D., Green, R. E., Kircher, M., Krause, J., Patterson, N., Durand, E. Y., et al. (2010). Genetic history of an archaic hominin group from Denisova Cave in Siberia. Nature, 468, 1053–1060.

    Article  Google Scholar 

  • Reich, D., Patterson, M., Kircher, M., Delfin, F., Nandineni, M. R., Pugach, I., et al. (2011). Denisova admixture and the first modern human dispersals into southeast Asia and Oceania. American Journal of Human Genetics, 89, 516–528.

    Article  Google Scholar 

  • Reynolds, T. (2013). The Middle Paleolithic of Cyrenaica: Is there an Aterian at the Haua Fteah and does it matter? Quaternary International, 300, 171–181.

    Article  Google Scholar 

  • Richter, D., Moser, J., Nami, M., Eiwanger, J., & Mikdad, A. (2010). New chronometric data from Ifri n’Ammar (Morocco) and the chronostratigraphy of the Middle Paleolithic in the Western Maghreb. Journal of Human Evolution, 59, 672–679.

    Article  Google Scholar 

  • Rightmire, G. P. (1976). Relationships of Middle and Upper Pleistocene hominids from sub-Saharan Africa. Nature, 260, 238–240.

    Article  Google Scholar 

  • Rightmire, G. P. (1978). Human skeletal remains from the southern Cape Province and their bearing on the Stone Age prehistory of South Africa. Quaternary Research, 9, 219–230.

    Article  Google Scholar 

  • Rightmire, G. P. (1979). Implications of the Border Cave skeletal remains for Later Pleistocene human evolution. Current Anthropology, 20, 23–35.

    Article  Google Scholar 

  • Rightmire, G. P. (1984). Homo sapiens in Sub-Saharan Africa. In F. H. Smith & R. Spencer (Eds.), The origins of modern humans: A world survey of the fossil evidence (pp. 295–325). New York: Alan R. Liss.

    Google Scholar 

  • Rightmire, G. P. (1998). Human evolution in the Middle Pleistocene: The role of Homo heidelbergensis. Evolutionary Anthropology, 6, 218–227.

    Article  Google Scholar 

  • Rightmire, G. P. (2001). Patterns of hominid evolution and dispersal in the Middle Pleistocene. Quaternary International, 75, 77–84.

    Article  Google Scholar 

  • Rightmire, G. P. (2008). Homo in the Middle Pleistocene: Hypodigms, variation, and species recognition. Evolutionary Anthropology, 17, 8–21.

    Article  Google Scholar 

  • Rightmire, G. P. (2009). Middle and later Pleistocene hominins in Africa and Southwest Asia. Proceedings of the National Academy of Sciences of the United States of America, 106, 16046–16050.

    Article  Google Scholar 

  • Rightmire, G. P., & Deacon, H. J. (1991). Comparative studies of Late Pleistocene human remains from Klasies River Mouth, South Africa. Journal of Human Evolution, 20, 131–156.

    Article  Google Scholar 

  • Rightmire, G. P., & Deacon, H. J. (2001). New human teeth from the Middle Stone Age deposits at Klasies River, South Africa. Journal of Human Evolution, 41, 535–544.

    Article  Google Scholar 

  • Rightmire, G. P., Deacon, H. J., Schwartz, J. H., & Tattersall, I. (2006). Human foot bones from Klasies River Main Site, South Africa. Journal of Human Evolution, 50, 96–103.

    Article  Google Scholar 

  • Roche, J. (1953). La grotte de Taforalt. L’Anthropologie, 57, 375–380.

    Google Scholar 

  • Roche, J. (1963). L’Épipaléolithique Marocaine. Lisbon: Fondation Calouste Gulbenkian.

    Google Scholar 

  • Roche, J. (1969). Les industries paléolithiques de la grotte de Taforalt (Maroc oriental). Quaternaria, 11, 89–100.

    Google Scholar 

  • Roche, J., & Texier, J. P. (1976). Découverte de restes humains dans un niveau Atérian de la grotte des Contrebandiers à Témara, Maroc. Comptes Rendus des Séances de l’Académie des Sciences (Paris), 282, 45–47.

    Google Scholar 

  • Rougier, H., Milota, Ş., Rodrigo, R., Gherase, M., Sarcină, L., Moldovan, O., et al. (2007). Peştera cu Oase 2 and the cranial morphology of early modern Europeans. Proceedings of the National Academy of Sciences of the United States of America, 104, 1165–1170.

    Article  Google Scholar 

  • Royer, D. F., Lockwood, C. A., Scott, J. E., & Grine, F. E. (2009). Size variation in early human mandibles and molars from Klasies River, South Africa: Comparison with other Middle and Late Pleistocene assemblages and with modern humans. American Journal of Physical Anthropology, 140, 312–323.

    Article  Google Scholar 

  • Saban, R. (1975). Les restes humains de Rabat (Kebibat). Annals de Paléontologie (Vertébrés), 61, 153–207.

    Google Scholar 

  • Saban, R. (1977). The place of Rabat Man (Kébibat, Morocco) in human evolution. Current Anthropology, 18, 518–524.

    Article  Google Scholar 

  • Salama, N., Guillemin, K., McDaniel, T. K., Sherlock, G., Tompkins, L., & Falkow, S. (2000). A whole-genome microarray reveals genetic diversity among Helicobacter pylori strains. Proceedings of the National Academy of Sciences of the United States of America, 97, 14668–14673.

    Article  Google Scholar 

  • Salas, A., Richards, M., De la Fe, T., Lareu, M. V., Sobrino, B., Sanchez-Diz, P., et al. (2002). The making of the African mtDNA landscape. American Journal of Human Genetics, 71, 1082–1111.

    Article  Google Scholar 

  • Salzmann, U., Hoelzmann, P., & Morczinek, I. (2002). Late Quaternary climate and vegetation of the Sudanian Zone of northeast Nigeria. Quaternary Research, 58, 73–83.

    Article  Google Scholar 

  • Sandford, K. S. (1934). Paleolithic man and the Nile Valley in Upper and Middle Egypt: A study of the region during Pliocene and Pleistocene times. Chicago: University of Chicago Press.

    Google Scholar 

  • Savage, D. K. (1983). Identifying industries in south central Africa. Ph.D. Dissertation, University of California, Berkeley.

    Google Scholar 

  • Sawchuk, E. A., & Willoughby, P. R. (2015). Terminal Pleistocene Later Stone Age human remains from the Mlambalasi Rock Shelter, Iringa Region, southern Tanzania. International Journal of Osteoarchaeology, 25, 593–607.

    Google Scholar 

  • Sawyer, S., Renauda, G., Violab, B., Hublin, J. J., Gansaugea, M. T., Shunkovd, M. V., et al. (2015). Nuclear and mitochondrial DNA sequences from two Denisovan individuals. Proceedings of the National Academy of Sciences of the United States of America, 112, 15696–15700.

    Google Scholar 

  • Scerri, E. M. L. (2013). The Aterian and its place in the North African Middle Stone Age. Quaternary International, 300, 111–130.

    Article  Google Scholar 

  • Schwartz, G. T., & Tattersall, I. (2000). The human chin revisited: What is it and who has it? Journal of Human Evolution, 38, 367–409.

    Article  Google Scholar 

  • Schefuß, E., Schouten, S., Jansen, J. H. F., & Damsté, J. S. S. (2003). African vegetation controlled by tropical sea surface temperatures in the mid-Pleistocene period. Nature, 422, 418–421.

    Article  Google Scholar 

  • Schefuß, E., Schouten, S., & Schneider, R. R. (2005). Climatic controls on central African hydrology during the past 20,000 years. Nature, 437, 1003–1006.

    Article  Google Scholar 

  • Schellmann, G., & Radtke, U. (1999). Problems encountered in the determination of dose and dose rate in ESR dating of mollusk shells. Quaternary Science Reviews, 18, 1515–1527.

    Article  Google Scholar 

  • Schlebusch, C. M., Skoglund, P., Sjodin, P., Gattepaille, L. M., Hernandez, D., Jay, F., et al. (2012). Genomic variation in seven Khoe-San groups reveals adaptation and complex African history. Science, 338, 374–379.

    Article  Google Scholar 

  • Schlebusch, C. M., Lombard, M., & Soodyall, H. (2013). mtDNA control region variation affirms diversity and deep sub-structure in populations from southern Africa. BMC Evolutionary Biology, 13, 56. doi:10.1186/1471-2148-13-56.

    Article  Google Scholar 

  • Scholz, C. A., Johnson, T. C., Cohen, A. S., King, J. W., Peck, J. A., Overpeck, J. T., et al. (2007). East African megadroughts between 135 and 75 thousand years ago and bearing on early-modern human origins. Proceedings of the National Academy of Sciences of the United States of America, 104, 16416–16421.

    Article  Google Scholar 

  • Scholz, C. A., Cohen, A. S., Johnson, T. C., King, J., Talbot, M. R., & Brown, E. T. (2011). Scientific drilling in the Great Rift Valley: The 2005 Lake Malawi Scientific Drilling Project—an overview of the past 145,000 years of climate variability in southern Hemisphere East Africa. Paleogeography, Paleoclimatology, Paleoecology, 303, 1–4.

    Article  Google Scholar 

  • Schultz, H., Emeis, K. C., Erlenkeuser, H., von Rad, U., & Rolf, C. (2002). The Toba volcanic event and interstadial/stadial climates at the Marine Isotopic Stage 5 to 4 transition in the northern Indian Ocean. Quaternary Research, 57, 22–31.

    Article  Google Scholar 

  • Schuster, S. C., Miller, W., Ratan, A., Tomsho, L. P., Giardine, B., Kasson, L. R., et al. (2010). Complete Khoisan and Bantu genomes from southern Africa. Nature, 463, 943–947.

    Article  Google Scholar 

  • Schwarcz, H., & Rink, J. (2000). ESR dating of the Die Kelders Cave 1 site, South Africa. Journal of Human Evolution, 38, 121–128.

    Article  Google Scholar 

  • Schwenninger, J. L., Collcutt, S. N., Barton, N., Bouzouggar, A., Clark Balzan, L., El Hajraoui, M. A., et al. (2010). A new luminescence chronology for Aterian cave sites on the Atlantic coast of Morocco. In E. A. Garcea (Ed.), South-eastern Mediterranean peoples between 130,000 and 10,000 years ago (pp. 18–36). Oxford: Oxbow Books.

    Google Scholar 

  • Scott, L. (1987). Pollen analysis of hyena coprolites and sediments from Equus Cave, Taung, Southern Kalahari (South Africa). Quaternary Research, 28, 144–156.

    Article  Google Scholar 

  • Scozzari, R., Cruciani, F., Santolamazza, P., Malaspina, P., Torroni, A., Sellitto, D., et al. (1999). Combined use of biallelic and microsatellite Y-chromosome polymorphisms to infer affinities among African populations. American Journal of Human Genetics, 65, 829–846.

    Article  Google Scholar 

  • Semino, O., Santachiara-Benerecetti, A. S., Falaschi, F., Cavalli-Sforza, L. L., & Underhill, P. A. (2002). Ethiopians and Khoisan share the deepest clades of the human Y-chromosome phylogeny. American Journal of Human Genetics, 70, 265–268.

    Article  Google Scholar 

  • Senut, B., Pickford, M., Braga, J., Marais, D., & Coppens, Y. (2000). Découverte d’un Homo sapiens archaïque à Oranjemund, Namibie. Comptes Rendus des Séances de l’Académie des Sciences (Paris) Sciences de la Terre et des Planètes, 330, 813–819.

    Google Scholar 

  • Şenyürek, M. S. (1940). Fossil Man in Tangier. Papers of the Peabody Museum of American Archaeology and Ethnology, Harvard University, 16, 1–27

    Google Scholar 

  • Shackelford, L. L. (2007). Regional variation in the postcranial robusticity of late Upper Paleolithic humans. American Journal of Physical Anthropology, 133, 655–668.

    Article  Google Scholar 

  • Shackleton, N. J. (1967). Oxygen isotope analyses and Pleistocene temperatures re-assessed. Nature, 215, 15–17.

    Article  Google Scholar 

  • Shackleton, N. J. (1969). The last interglacial in the marine and terrestrial records. Proceedings of the Royal Society B, 174, 135–154.

    Article  Google Scholar 

  • Shackleton, N. J. (1982). Stratigraphy and chronology of the Klasies River Mouth deposits: Oxygen isotope evidence. In R. Singer & J. Wymer (Eds.), The Middle Stone Age at Klasies River Mouth in South Africa (pp. 194–199). Chicago: University of Chicago Press.

    Google Scholar 

  • Shackleton, N. J. (1987). Oxygen isotopes, ice volume and sea level. Quaternary Science Reviews, 6, 183–190.

    Article  Google Scholar 

  • Shackleton, N. J., & Pisias, N. G. (1985). Atmospheric carbon dioxide, orbital forcing, and climate. In E. Sundquist & W. S. Broecker (Eds.), The carbon cycle and atmospheric CO 2 : Natural variations Archean to present (Geophysical Monograph Series No. 32) (pp. 303–317). Washington, DC: American Geophysical Union.

    Google Scholar 

  • Shackleton, N. J., Berger, A., & Peltier, W. R. (1990). An alternative astronomical calibration of the Lower Pleistocene time scale based on ODP Site 677. Transactions of the Royal Society of Edinburgh, Earth Science, 81, 251–261.

    Article  Google Scholar 

  • Shaw, T., & Daniels, S. G. H. (1984). Excavations at Iwo Eleru, Ondo State, Nigeria. West African Journal of Archaeology, 14, 1–269.

    Article  Google Scholar 

  • Shea, J. J., Fleagle, J. G., & Assefa, Z. (2007). Context and chronology of early Homo sapiens fossils from the Omo Kibish Formation, Ethiopia. In P. Mellars, K. Boyle, O. Bar-Yosef, & C. Stringer (Eds.), Rethinking the human revolution (pp. 153–162). Cambridge: McDonald Institute for Archaeological Research.

    Google Scholar 

  • Sillen, A., & Morris, A. (1996). Diagenesis of bone from Border Cave: Implications for the age of the Border Cave hominids. Journal of Human Evolution, 31, 499–506.

    Article  Google Scholar 

  • Singer, R. (1993). Mythical African ‘Australoids’ and triangular bricks: The Cape Flats skull in retrospect. South African Archaeological Bulletin, 48, 105–112.

    Article  Google Scholar 

  • Singer, R., & Wymer, J. J. (1982). The Middle Stone Age at Klasies River Mouth in South Africa. Chicago: University of Chicago Press.

    Google Scholar 

  • Smith, F. H. (1985). Continuity and change in the origin of modern Homo sapiens. Zeitschrift für Morphologie und Anthropologie, 75, 197–222.

    Google Scholar 

  • Smith, F. H. (1992). Models and realities in modern human origins: The African fossil evidence. Philosophical Transactions of the Royal Society B, 337, 243–250.

    Article  Google Scholar 

  • Smith, F. H. (2002). Migrations, radiations and continuity: Patterns in the evolution of Middle and Late Pleistocene humans. In W. C. Hartwig (Ed.), The primate fossil record (pp. 437–456). Cambridge: Cambridge University Press.

    Google Scholar 

  • Smith, F., Falsetti, A., & Donnelly, S. (1989). Modern human origins. Yearbook of Physical Anthropology, 32, 35–68.

    Article  Google Scholar 

  • Smith, P., & Shegev, M. (1988). The dentition of Nubians from Wadi Halfa, Sudan: An evolutionary perspective. Journal of the Dental Association of South Africa, 43, 535–541.

    Google Scholar 

  • Smith, P., Nshimirimana, R., de Beer, F. Morris, D., Jacobson, L., Chazan, M., et al. (2012). Canteen Kopje: A new look at an old skull. South African Journal of Science, 108(1/2), Article 738, 9 pages. doi:10.4102/sajs.v108il/2.738.

  • Smith, T. M., Olejniczak, A. J., Tafforeau, P., Reid, D. J., Grine, F. E., & Hublin, J. J. (2006). Molar crown thickness, volume, and development in South African Middle Stone Age humans. South African Journal of Science, 102, 513–517.

    Google Scholar 

  • Smith, T., Tafforeau, P., Reid, D., Grün, R., Eggins, S., Boutakiout, M., et al. (2007). Earliest evidence of modern human life history in North African early Homo sapiens. Proceedings of the National Academy of Sciences of the United States of America, 104, 6128–6133.

    Article  Google Scholar 

  • Soficaru, A., Doboş, A., & Trinkaus, E. (2006). Early modern humans from the Peştera Muierii, Baia de Fier, Romania. Proceedings of the National Academy of Sciences of the United States of America, 103, 17196–17201.

    Article  Google Scholar 

  • Soficaru, A., Petrea, C., Doboş, A., & Trinkaus, E. (2007). The human cranium from Peştera Cioclovina Uscată, Romania. Context, age, taphonomy, morphology, and paleopathology. Current Anthropology, 48, 611–619.

    Article  Google Scholar 

  • Spoor, F., & Stringer, C. (1998). Rare temporal bone pathology of the Singa calvaria from Sudan. American Journal of Physical Anthropology, 107, 41–50.

    Article  Google Scholar 

  • Spoor, F., Leakey, M. G., Antón, S. C., & Leakey, L. N. (2008). The taxonomic status of KNM-ER 42700: A reply to Baab (2008a). Journal of Human Evolution, 55, 747–750.

    Article  Google Scholar 

  • Starling, A. P., & Stock, J. T. (2007). Dental indicators of health and stress in early Egyptian and Nubian agriculturalists: A difficult transition and gradual recovery. American Journal of Physical Anthropology, 134, 520–528.

    Article  Google Scholar 

  • Stearns, C. E., & Thurber, D. L. (1965). Th230/U234 dates of Late Pleistocene marine fossils from the Mediterranean and Moroccan littorals. Progress in Oceanography, 4, 293–305.

    Article  Google Scholar 

  • Stock, J. T., O’Neill, M. C., Ruff, C. B., Zabecki, M., Shackelford, L., & Rose, J. C. (2011). Body size, skeletal biomechanics, mobility and habitual activity from the Late Paleolithic to the Mid-Dynastic Nile Valley. In R. Pinhasi & J. T. Stock (Eds.), Human bioarchaeology of the transition to agriculture (pp. 347–369). New York: Wiley.

    Chapter  Google Scholar 

  • Stocker, T. F. (1998). Climate change: The seesaw effect. Science, 282, 61–62.

    Article  Google Scholar 

  • Stokes, S., & Bailey, R. (2002). Optical dating of Nazlet Safaha and Nazlet Khater 4. In P. M. Vermeersch (Ed.), Paleolithic quarrying sites in upper and middle Egypt (pp. 349–351). Leuven: Leuven University Press.

    Google Scholar 

  • Stokes, S., Thomas, D. S. G., & Washington, R. (1997). Multiple episodes of aridity in southern Africa since the last interglacial period. Nature, 388, 154–158.

    Article  Google Scholar 

  • Stringer, C. B. (1974). Population relationships of later Pleistocene hominids: A multivariate study of available crania. Journal of Archaeological Science, 1, 317–342.

    Article  Google Scholar 

  • Stringer, C. B. (1978). Some problems in Middle and Upper Pleistocene hominid relationships. In D. Chivers & K. Joysey (Eds.), Recent advances in primatology (Vol. 3, pp. 395–418)., evolution London: Academic.

    Google Scholar 

  • Stringer, C. B. (2002). Modern human origins: Progress and prospects. Philosophical Transactions of the Royal Society London B, 357, 563–579.

    Article  Google Scholar 

  • Stringer, C. B. (2007). The origin and dispersal of Homo sapiens: Our current state of knowledge. In P. Mellars, K. Boyle, O. Bar-Yosef, & C. Stringer (Eds.), Rethinking the human revolution (pp. 15–20). Cambridge: McDonald Institute for Archaeological Research.

    Google Scholar 

  • Stringer, C. B., & Bräuer, G. (1994). Methods, misreading, and bias. American Anthropologist, 96, 416–424.

    Article  Google Scholar 

  • Stringer, C. B., & Andrews, P. (1988). Genetic and fossil evidence for the origin of modern humans. Science, 239, 1263–1268.

    Article  Google Scholar 

  • Stringer, C. B., Cornish, L., & Stuart-Macadam, P. (1985). Preparation and further study of the Singa skull from Sudan. Bulletin of the British Museum Natural History (Geology), 38, 347–358.

    Google Scholar 

  • Stynder, D. D., Ackermann, R. R., & Sealy, J. C. (2007a). Craniofacial variation and population continuity during the South African Holocene. American Journal of Physical Anthropology, 134, 489–500.

    Article  Google Scholar 

  • Stynder, D. D., Ackermann, R. R., & Sealy, J. C. (2007b). Early to mid-Holocene South African Later Stone Age human crania exhibit a distinctly Khoesan morphological pattern. South African Journal of Science, 103, 349–352.

    Google Scholar 

  • Stynder, D., Brock, F., Sealy, J., Wurz, S., Morris, A., & Volman, T. (2009). A mid-Holocene AMS 14C date for the presumed Upper Pleistocene human skeleton from Peers Cave, South Africa. Journal of Human Evolution, 56, 431–434.

    Article  Google Scholar 

  • Tang, H., Siegmund, D. O., Shen, P., Oefner, P. J., & Feldman, M. W. (2002). Frequentist estimation of coalescence times from nucleotide sequence data using a tree-based partition. Genetics, 161, 447–459.

    Google Scholar 

  • Tattersall, I., & Schwartz, J. H. (1999). Hominids and hybrids: The place of Neanderthals in human evolution. Proceedings of the National Academy of Sciences of the United States of America, 96, 7117–7119.

    Article  Google Scholar 

  • Teilhard de Chardin, P. (1930). Le Paléolithique en Somalie francaise et en Abyssinie. L’ Anthropologie, 40, 331–334.

    Google Scholar 

  • Texier, P. J., Porraz, G., Parkington, J., Rigaud, J.-P., Poggenpoel, C., Miller, C., et al. (2010). A Howiesons Poort tradition of engraving ostrich eggshell containers dated to 60,000 years ago at Diepkloof Rock Shelter, South Africa. Proceedings of the National Academy of Science of the United States of America, 107, 6180–6185.

    Article  Google Scholar 

  • Thoma, A. (1984). Morphology and affinities of the Nazlet Khater man. Journal of Human Evolution, 13, 287–296.

    Article  Google Scholar 

  • Thoma, A., & Vallois, H. V. (1977). Les dents de l’homme de Rabat. Bulletins et Mémoires de la Societé d’Anthropologie de Paris, 4, 31–58.

    Article  Google Scholar 

  • Thomas, D. S. G., & Burrough, S. L. (2012). Interpreting geoproxies of late Quaternary climate change in African drylands: Implications for understanding environmental change and early human behavior. Quaternary International, 253, 5–17.

    Article  Google Scholar 

  • Thomas, D. S. G., & Shaw, P. A. (2002). Late Quaternary environmental change in central southern Africa: New data, synthesis, issues and prospects. Quaternary Science Reviews, 21, 783–797.

    Article  Google Scholar 

  • Thomas, D. S. G., Bailey, R., Shaw, P. A., Durcan, J. A., & Singarayer, J. S. (2009). Late Quaternary highstands at Lake Chilwa, Malawi: Frequency, timing and possible forcing mechanisms in the last 44 ka. Quaternary Science Reviews, 28, 526–539.

    Article  Google Scholar 

  • Thompson, W. G., & Goldstein, S. L. (2006). A radiometric calibration of the SPECMAP timescale. Quaternary Science Reviews, 25, 3207–3215.

    Article  Google Scholar 

  • Thomson, R., Pritchard, J. K., Shen, P., Oefner, P. J., & Feldman, M. W. (2000). Recent common ancestry of human Y chromosomes: Evidence from DNA sequence data. Proceedings of the National Academy of Sciences of the United States of America, 97, 7360–7365.

    Article  Google Scholar 

  • Tierney, J. E., & deMenocal, P. B. (2013). Abrupt shifts in Horn of Africa hydroclimate since the Last Glacial Maximum. Science, 342, 843–846.

    Article  Google Scholar 

  • Tierney, J. E., Russell, J. M., Huang, Y., Damsté, J. S. S., Hopmans, E. C., & Cohen, A. S. (2008). Northern hemisphere controls on tropical southeast African climate during the past 60,000 years. Science, 322, 252–255.

    Article  Google Scholar 

  • Tierney, J. E., Lewis, S. C., Cook, B. I., LeGrande, A. N., & Schmidt, G. A. (2011). Model, proxy and isotopic perspectives on the East African Humid Period. Earth and Planetary Science Letters, 307, 103–112.

    Article  Google Scholar 

  • Tishkoff, S. A., & Verrelli, B. C. (2003). Role of evolutionary history on haplotype block structure in the human genome: Implications for disease mapping. Current Opinion in Genetics & Development, 13, 569–575.

    Article  Google Scholar 

  • Tishkoff, S. A., Gonder, M. K., Henn, B. M., Mortensen, H., Knight, A., Gignoux, C., et al. (2007). History of click-speaking populations of Africa inferred from mtDNA and Y chromosome genetic variation. Molecular Biology and Evolution, 24, 2180–2195.

    Article  Google Scholar 

  • Tishkoff, S. A., Reed, F. A., Friedlaender, F. R., Ehret, C., Ranciaro, A., Froment, A., et al. (2009). The genetic structure and history of Africans and African Americans. Science, 324, 1035–1044.

    Article  Google Scholar 

  • Tobias, P. V. (1967). The hominid skeletal remains of Haua Fteah. Appendix 1B. In C. B. M. McBurney (Ed.), The Haua Fteah (Cyrenaica) and the Stone Age of the south-east Mediterranean (pp. 338–352). Cambridge: Cambridge University Press.

    Google Scholar 

  • Toerien, M. J., & Hughes, A. R. (1955). The limb bones of Springbok Flats Man. South African Journal of Science, 52, 125–128.

    Google Scholar 

  • Torroni, A., Achilli, A., Macaulay, V., Richards, M., & Bandelt, H. J. (2006). Harvesting the fruit of the human mtDNA tree. Trends in Genetics, 22, 339–345.

    Article  Google Scholar 

  • Trauth, M. H., Deino, A. L., Bergner, A. G. N., & Strecker, M. R. (2003). East African climate change and orbital forcing during the last 175 kyr BP. Earth and Planetary Science Letters, 206, 297–313.

    Article  Google Scholar 

  • Trauth, M. H., Maslin, M. A., Deino, A., & Strecker, M. R. (2005). Late Cenozoic moisture history of East Africa. Science, 309, 2051–2053.

    Article  Google Scholar 

  • Trauth, M. H., Maslin, M. A., Deino, A., Bergner, A. G. N., Dühnforth, M., & Strecker, M. R. (2007). High- and low-latitude forcing of Plio-Pleistocene East African climate and human evolution. Journal of Human Evolution, 53, 475–486.

    Article  Google Scholar 

  • Tribolo, C., Mercier, N., Selo, M., Joron, J. L., Reyss, J. L., Henshilwood, C., et al. (2006). TL dating of burnt lithics from Blombos Cave (South Africa): Further evidence for the antiquity of modern human behaviour. Archaeometry, 48, 341–357.

    Article  Google Scholar 

  • Tribolo, C., Mercier, N., Valladas, H., Joron, J. L., Guibert, P., Lefrais, Y., et al. (2009). Thermoluminescence dating of a Stillbay-Howiesons Poort sequence at Diepkloof Rock Shelter (Western Cape, South Africa). Journal of Archaeological Science, 36, 730–739.

    Article  Google Scholar 

  • Tribolo, C., Mercier, N., Douville, E., Joron, J. L., Reyss, J. L., Rufer, D., et al. (2013). OSL and TL dating of the Middle Stone Age sequence at Diepkloof Rock Shelter (South Africa): A clarification. Journal of Archaeological Science, 40, 3401–3411.

    Article  Google Scholar 

  • Trinkaus, E. (1993). A note on the KNM-ER 999 hominid femur. Journal of Human Evolution, 24, 493–504.

    Article  Google Scholar 

  • Trinkaus, E. (2005). Early modern humans. Annual Review of Anthropology, 34, 207–230.

    Article  Google Scholar 

  • Trinkaus, E., Moldovan, O., Milota, Ş., Bîlgăr, A., Sarcina, L., Athreya, S., et al. (2003). An early modern human from the Peştera cu Oase, Romania. Proceedings of the National Academy of Sciences of the United States of America, 100, 11231–11236.

    Article  Google Scholar 

  • Tryon, C. A., Faith, J. T., Peppe, D. J., Fox, D. L., McNulty, K. P., Jenkins, K., et al. (2010). The Pleistocene archaeology and environments of the Wasiriya Beds, Rusinga Island, Kenya. Journal of Human Evolution, 59, 657–671.

    Article  Google Scholar 

  • Tryon, C. A., Peppe, D. J., Faith, J. T., Van Plantinga, A., Nightengale, S., Ogondo, J., et al. (2012). Late Pleistocene artefacts and fauna from Rusinga and Mfangano islands, Lake Victoria, Kenya. Azania: Archaeological Research in Africa, 47, 14–38.

    Article  Google Scholar 

  • Tryon, C. A., Crevecoeur, I., Faith, J. T., Ekshtain, R., Nivens, J., Patterson, D., et al. (2015). Late Pleistocene age and archaeological context for the hominin calvaria from GvJm-22 (Lukenya Hill, Kenya). Proceedings of the National Academy of Sciences of the United States of America, 112, 2682–2687.

    Article  Google Scholar 

  • Turner, C. G., & Markowitz, M. A. (1990). Dental discontinuity between Late Pleistocene and recent Nubians: Peopling of the Eurafrican-south Asian triangle, I. Homo, 41, 32–41.

    Google Scholar 

  • Twiesselmann, F. (1958). Les ossements humains du gîte mésolithique d’Ishango. Exploration du Parc national Albert, mission J. de Heinzelin de Braucourt, 1950 (Institut des Parcs nationaux du Congo belge, Fasc. 5). Brussels: Institut des Parcs nationaux du Congo belge.

    Google Scholar 

  • Tzedakis, C. (2003). Timing and duration of Last Interglacial conditions in Europe: A chronicle of a changing chronology. Quaternary Science Reviews, 22, 763–768.

    Article  Google Scholar 

  • Tzedakis, P. C., Andrieu, V., de Beaulieu, J.-L., Crowhurst, S., Follieri, M., Hooghiemstra, H., et al. (1997). Comparison of terrestrial and marine records of changing climate of the last 500,000 years. Earth and Planetary Science Letters, 150, 171–176.

    Article  Google Scholar 

  • Underhill, P. A., Shen, P., Lin, A. A., Jin, L., Passarino, G., Yang, W. H., et al. (2000). Y chromosome sequence variation and the history of human populations. Nature Genetics, 26, 358–361.

    Article  Google Scholar 

  • Underhill, P. A., Passarino, G., Lin, A. A., Shen, P., Lahr, M. M., Foley, R. A., et al. (2001). The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations. Annals of Human Genetics, 65, 43–62.

    Article  Google Scholar 

  • Urey, H. C. (1947). The thermodynamic properties of isotopic substances. Journal of the Chemical Society, 1947, 562.

    Article  Google Scholar 

  • Vallois, H. V. (1945). L’homme fossile de Rabat. Comptes Rendus à l’Académie des Sciences, 221, 669–671.

    Google Scholar 

  • Vallois, H. V. (1951). La mandibule humaine fossile de la grotte du Porc-epic près Dire Daoua (Abyssinie). L’ Anthropologie, 55, 231–238.

    Google Scholar 

  • Vallois, H. V. (1969). Les hommes de Cro-Magnon et les Guanches: Les faits acquis et les hypothéses. Simposio del Cro-Magnon. Anuario Estudias Atlanticos, Madrid, 15, 97–119.

    Google Scholar 

  • Vallios, H. V., & Roche, J. (1958). La mandibule Acheuléenne de Témara. Comptes Rendus des Séances de l’Académie des Sciences (Paris), 246, 3113–3116.

    Google Scholar 

  • van Andel, T. H., & Tzedakis, P. C. (1996). Paleolithic landscapes of Europe and environs 150,000–25,000 years ago: An overview. Quaternary Science Reviews, 15, 481–500.

    Article  Google Scholar 

  • Van Couvering, J. (1972). Geology of Rusinga Island and correlation of the Kenya mid-Tertiary fauna. Ph.D. Dissertation, University of Cambridge.

    Google Scholar 

  • Van Peer, P., Vermeersch, P. M., & Paulissen, E. (2010). Chert quarrying, lithic technology, and a modern human burial at the paleolithic site of Taramsa 1, upper Egypt. Leuven: Leuven University Press.

    Google Scholar 

  • van Riet Lowe, C. (1929). Notes on some stone implements from Tuinplaats, Springbok Flats. South African Journal of Science, 26, 623–630.

    Google Scholar 

  • van Riet Lowe, C. (1954). An artefact recovered with the Boskop calvaria. South African Archaeological Bulletin, 9, 135–137.

    Article  Google Scholar 

  • van Vark, G. N. (1984). On the determination of hominid affinities. In G. N. van Vark & W. W. Howells (Eds.), Multivariate statistical methods in physical anthropology (pp. 323–349). Doordrecht: D. Reidel.

    Chapter  Google Scholar 

  • van Vark, G. N., Bilsborough, A., & Dijkema, J. (1989). A further study of the morphological affinities of the Border Cave 1 cranium, with special reference to the origin of modern man. Anthropologie et Préhistoire, 100, 43–56.

    Google Scholar 

  • Veeramah, K. R., Wegmann, D., Woerner, A., Mendez, F. L., Watkins, J. C., Destro-Bisol, G., et al. (2012). An early divergence of KhoeSan ancestors from those of other modern humans is supported by an ABC-based analysis of autosomal resequencing data. Molecular Biology and Evolution, 29, 617–630.

    Article  Google Scholar 

  • Vermeersch, P. M. (2002). Two Upper Paleolithic burials at Nazlet Khater. In P. M. Vermeersch (Ed.), Paleolithic quarrying sites in upper and middle Egypt (pp. 273–282). Leuven: Leuven University Press.

    Google Scholar 

  • Vermeersch, P. M., Gijselings, G., & Paulissen, E. (1984). Discovery of the Nazlet Khater Man, upper Egypt. Journal of Human Evolution, 13, 281–286.

    Article  Google Scholar 

  • Vermeersch, P. M., Paulissen, E., Stokes, S., de Van Peer, P., Bie, M., Steenhoudt, F., & Missotten, S. (1997). Middle Paleolithic chert mining in Eqypt. In A. Ramos-Millán & M. A. Bustillo (Eds.), Siliceous rocks and culture (pp. 173–193). Granada, Spain: University of Granada.

    Google Scholar 

  • Vermeersch, P. M., Paulissen, E., Stokes, S., Charlier, C., Van Peer, P., Stringer, C., et al. (1998). A Middle Paleolithic burial of a modern human at Taramsa Hill, Egypt. Antiquity, 72, 475–484.

    Google Scholar 

  • Verna, C., Texier, P. J., Rigaud, J. P., Poggenpoel, C., & Parkington, J. (2013). The Middle Stone Age human remains from Diepkloof Rock Shelter (Western Cape, South Africa). Journal of Archaeological Science, 40, 3532–3541.

    Article  Google Scholar 

  • Vignard, E. (1928). Une nouvelle industrie lithique, le Sébilien. Bulletin de la Société Préhistorique Française, 25, 200–220.

    Article  Google Scholar 

  • Villa, P., Soriano, S., Teyssandier, N., & Wurz, S. (2010). The Howiesons Poort and MSA III at Klasies River Main Site, Cave 1A. Journal of Archaeological Science, 37, 630–655.

    Article  Google Scholar 

  • Villa, P., Soriano, S., Tsanova, T., Degano, I., Higham, T. F. G., d’Errico, F., et al. (2012). Border Cave and the beginning of the Later Stone Age in South Africa. Proceedings of the National Academy of Sciences of the United States of America, 109, 13208–13213.

    Article  Google Scholar 

  • Voelker, A. H. L., & Participants, Workshop. (2002). Global distribution of centennial-scale records for Marine Isotope Stage (MIS) 3: A database. Quaternary Science Reviews, 21, 1185–1212.

    Article  Google Scholar 

  • Vogel, J. C. (2001). Radiometric dates for the Middle Stone Age in South Africa. In P. V. Tobias, M. A. Raath, J. Moggi-Cecchi, & G. A. Doyle (Eds.), Humanity from African naissance to coming millennia-colloquia in human biology and paleoanthropology (pp. 261–268). Firenze: Firenza University Press.

    Google Scholar 

  • Vogel, J. C., & Beaumont, P. (1972). Revised radiocarbon chronology for the Stone Age in South Africa. Nature, 237, 50–51.

    Article  Google Scholar 

  • Vogel, J. C., & Waterbolk, H. T. (1963). Groningen radiocarbon dates IV. Radiocarbon, 5, 163–202.

    Google Scholar 

  • Vogel, J. C., Fuls, A., & Visser, E. (1986). Pretoria radiocarbon dates III. Radiocarbon, 28, 1133–1172.

    Google Scholar 

  • Volman, T. P. (1984). Early prehistory of southern Africa. In R. G. Klein (Ed.), Southern African prehistory and paleoenvironments (pp. 169–220). Rotterdam: Balkema.

    Google Scholar 

  • Wadley, L. (1993). The Pleistocene Later Stone Age south of the Limpopo River. Journal of World Prehistory, 7, 243–296.

    Google Scholar 

  • Wadley, L. (2007). Announcing a Still Bay industry at Sibudu Cave, South Africa. Journal of Human Evolution, 52, 681–689.

    Article  Google Scholar 

  • Wadley, L., & Jacobs, Z. (2004). Sibudu Cave, KwaZulu-Natal: Background to the excavations of Middle Stone Age and Iron Age occupations. South African Journal of Science, 100, 145–151.

    Google Scholar 

  • Wadley, L., & Jacobs, Z. (2006). Sibudu cave: Background to the excavations, stratigraphy and dating. Southern African Humanities, 18, 1–26.

    Google Scholar 

  • Wadley, L., Hodgskiss, T., & Grant, M. (2009). Implications for complex cognition from the hafting of tools with compound adhesives in the Middle Stone Age, South Africa. Proceedings of the National Academy of Sciences of the United States of America, 106, 9590–9594.

    Article  Google Scholar 

  • Wadley, L., Sievers, C., Bamford, M., Goldberg, P., Berna, F., & Miller, C. (2011). Middle Stone Age bedding construction and settlement patterns at Sibudu, South Africa. Science, 334, 1388–1391.

    Article  Google Scholar 

  • Wall, J. D., Lohmueller, K. E., & Plagnol, V. (2009). Detecting ancient admixture and estimating demographic parameters in multiple human populations. Molecular Biology and Evolution, 26, 1823–1827.

    Article  Google Scholar 

  • Wall, J. D., Yang, M. A., Jay, F., Kim, S. K., Durand, E. Y., Stevinson, L. S., et al. (2013). Higher levels of Neanderthal ancestry in East Asians than in Europeans. Genetics, 194, 199–209.

    Article  Google Scholar 

  • Wang, Q., Tobias, P., Roberts, D., & Jacobs, Z. (2008). A re-examination of a human femur found at the Blind River site, East London, South Africa: Its age, morphology, and breakage pattern. Anthropological Review, 71, 43–61.

    Article  Google Scholar 

  • Watts, I. (2010). The pigments from Pinnacle Point Cave 13B, Western Cape, South Africa. Journal of Human Evolution, 59, 392–411.

    Article  Google Scholar 

  • Watts, W. A., Allen, J. R. M., & Huntley, B. (2000). Paleoecology of three interstadial events during oxygen-isotope Stages 3 and 4: A lacustrine record from Lago Grande di Monticchio, southern Italy. Paleogeography, Paleoclimatology, Paleoecology, 155, 83–93.

    Article  Google Scholar 

  • Weaver, T. D. (2012). Did a discrete event 200,000-100,000 years ago produce modern humans? Journal of Human Evolution, 63, 121–126.

    Article  Google Scholar 

  • Weldeab, S., Schneider, R. R., Kölling, M., & Wefer, G. (2005). Holocene African droughts relate to eastern equatorial Atlantic cooling. Geology, 33, 981–984.

    Article  Google Scholar 

  • Wells, L. H. (1935). A fossilized human femur from East London, Cape Province. South African Journal of Science, 32, 596–600.

    Google Scholar 

  • Wells, L. H. (1951). The fossil human skull from Singa. Fossil mammals of Africa (pp. 29–42). London: British Museum of Natural History.

    Google Scholar 

  • Wells, L H. (1952). Human crania of the Middle Stone Age in South Africa. Proceedings of the first pan-African congress on prehistory, Nairobi, 1947 (pp. 125–133). Oxford: Blackwell.

    Google Scholar 

  • Wells, L. H. (1959). The problem of Middle Stone Age Man in southern Africa. Man, 244, 158–160.

    Article  Google Scholar 

  • Wendorf, F. (1968). Site 117: A Nubian final Paleolithic graveyard near Jebel Sahaba, Sudan. In F. Wendorf (Ed.), The Prehistory of Nubia (Vol. 2, pp. 954–995). Dallas: Southern Methodist University Press.

    Google Scholar 

  • Wendorf, F., & Schild, R. (1976). Prehistory of the Nile Valley. New York: Academic.

    Google Scholar 

  • Wendorf, F., & Schild, R. (2004). Late Paleolithic warfare in Nubia: The evidence and causes. Adumatu (Journal of the Arab World), 10, 7–28.

    Google Scholar 

  • Wendorf, F., Said, S., & Schild, R. (1970). Egyptian prehistory: Some new concepts. Science, 169, 1161–1171.

    Article  Google Scholar 

  • Wendorf, F., Schild, R., & Haas, H. (1979). A new radiocarbon chronology for prehistoric sites in Nubia. Journal of Field Archaeology, 6, 219–223.

    Google Scholar 

  • White, T. D., Asfaw, B., DeGusta, D., Gilbert, H., Richards, G. D., Suwa, G., et al. (2003). Pleistocene Homo sapiens from Middle Awash, Ethiopia. Nature, 423, 742–747.

    Article  Google Scholar 

  • Williams, D. F., Thunell, R. C., Tappa, E., Rio, D., & Raffi, I. (1988). Chronology of the Pleistocene oxygen isotope record: 0–1.88 m.y. B.P. Paleogeography, Paleoclimatology, Paleoecology, 64, 221–240.

    Article  Google Scholar 

  • Willoughby, P. R. (2012). The Middle and Later Stone Age in the Iringa Region of southern Tanzania. Quaternary International, 270, 103–118.

    Article  Google Scholar 

  • Wolpoff, M. H., Frayer, D. W., & Jelínek, J. (2006). Aurignacian female crania and teeth from the Mladeč caves, Moravia, Czech Republic. In M. Teschler-Nicola (Ed.), Early modern humans at the Moravian Gate: The Mladeč caves and their remains (pp. 273–340). Vienna: Springer.

    Chapter  Google Scholar 

  • Woodward, A. S. (1938). A fossil skull of an ancestral Bushman from the Anglo-Egyptian Sudan. Antiquity, 12, 193–195.

    Google Scholar 

  • Wright, J. D. (2000). Global climate change in marine stable isotope records. In J. S. Noller, J. M. Sowers, & W. R. Lettis (Eds.), Quaternary geology methods and applications (pp. 427–433). Washington, DC: American Geophysical Union.

    Chapter  Google Scholar 

  • Wrinn, P. J., & Rink, W. J. (2003). ESR dating of tooth enamel from Aterian levels at Mugharet el’Aliya (Tangier, Morocco). Journal of Archaeological Science, 30, 123–133.

    Article  Google Scholar 

  • Wurz, S. (1999). The Howiesons Poort backed artefacts from Klasies River: An argument for symbolic behavior. South African Archaeology Bulletin, 54, 38–50.

    Article  Google Scholar 

  • Wurz, S. (2002). Variability in the Middle Stone Age lithic sequence, 115,000–60,000 years ago at Klasies River, South Africa. Journal of Archaeological Science, 29, 1001–1015.

    Article  Google Scholar 

  • Wurz, S. (2008). Modern behavior at Klasies River. South African Archaeological Society Goodwin Series, 10, 150–156.

    Google Scholar 

  • Wurz, S. (2012). The significance of MIS 5 shell middens on the Cape coast: A lithic perspective from Klasies River and Ysterfontein 1. Quaternary International, 270, 61–69.

    Article  Google Scholar 

  • Wymer, J. (1982). The paleolithic age. New York: St. Martin’s Press.

    Google Scholar 

  • Wynn, J. G. (2006). Influence of Plio-Pleistocene aridification on human evolution: Evidence from paleosols of the Turkana Basin, Kenya. American Journal of Physical Anthropology, 123, 106–118.

    Article  Google Scholar 

  • Yellen, J., Brooks, A., Helgren, D., Tappen, M., Ambrose, S., Bonnefille, R., et al. (2005). The archaeology of Aduma Middle Stone Age sites in the Awash Valley, Ethiopia. PaleoAnthropology, 3, 25–100.

    Google Scholar 

  • Yokoyama, Y., Falguères, C., Sémah, F., Jacob, T., & Grün, R. (2008). Gamma-ray spectrometric dating of late Homo erectus skulls from Ngandong and Sambungmacan, central Java, Indonesia. Journal of Human Evolution, 55, 274–277.

    Article  Google Scholar 

  • Zhao, J. X., Hu, K., Collerson, K. D., & Xu, H. K. (2001). Thermal ionization mass spectrometry U-series dating of a hominid site near Nanjing, China. Geology, 29, 27–30.

    Article  Google Scholar 

  • Zhao, M., Dupont, L., Eglington, G., & Teece, M. (2003). n-Alkane and pollen reconstruction of terrestrial climate and vegetation for N.W. Africa over the last 160 kyr. Organic Geochemistry, 34, 131–143.

    Article  Google Scholar 

  • Ziegler, M., Simon, M. H., Hall, I. R., Barker, S., Stringer, S., & Zahn, R. (2013). Development of Middle Stone Age innovation linked to rapid climate change. Nature Communications, 4, 1905. doi:10.1038/ncomms2897.

    Article  Google Scholar 

  • Zielinski, G. A., Mayewski, P. A., Meeker, L. D., Whitlow, S., Twickler, M. S., & Taylor, K. (1996). Potential atmospheric impact of the Toba mega-eruption 71,000 years ago. Geophysical Research Letters, 23, 837–840.

    Article  Google Scholar 

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Acknowledgements

I thank Brian Stewart for the invitation to provide this review, and Brian, Sacha Jones and Eric Delson for their persistence and patience. Thanks to Zelalem Assefa, Graeme Barker, Frank Brown, Isabelle Crevecoeur, Tyler Faith, Craig Feibel, John Fleagle, Chris Henshilwood, Jean-Jacques Hublin, Chris Hunt, Richard Klein, Christine Lane, Curtis Marean, Justin Pargeter, John Parkington, Tim Reynolds, Eleanor Scerri, John Shea, Lyn Wadley, Tim White, Pamela Willoughby and Sarah Wurz for information pertaining to specific Late Pleistocene sites and fossils. I am grateful to Jenna Cole for information and discussions concerning paleoclimate and late Quaternary African paleoenvironments, and to Brenna Henn for providing invaluable insights into the human genetic landscape of Africa. I am especially grateful to Eric Delson, John Fleagle, Tim Compton, Chris Stringer and Richard Klein for their comments and suggestions on the entire manuscript. The illustrations were skillfully executed by Luci Betti-Nash.

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Grine, F.E. (2016). The Late Quaternary Hominins of Africa: The Skeletal Evidence from MIS 6-2. In: Jones, S., Stewart, B. (eds) Africa from MIS 6-2. Vertebrate Paleobiology and Paleoanthropology. Springer, Dordrecht. https://doi.org/10.1007/978-94-017-7520-5_17

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