Skip to main content
Log in

Increasing Behavioral Flexibility? An Integrative Macro-Scale Approach to Understanding the Middle Stone Age of Southern Africa

  • Published:
Journal of Archaeological Method and Theory Aims and scope Submit manuscript

Abstract

The Middle Stone Age (MSA) of southern Africa represents a period during which anatomically modern humans adopted a series of diverse cultural innovations. Researchers generally attribute these behavioral changes to environmental, neurological, or demographic causes, but none of these alone offers a satisfactory explanation. Even as patterns at site level come into focus, large-scale trends in cultural expansions remain poorly understood. This paper presents different ways to view diachronic datasets from localities in southern Africa and specifically tests hypotheses of environmental and cultural causality. We employ an array of analyses in an attempt to understand large-scale variability observed during the MSA. We evaluated the diversity of stone tool assemblages to model site use, examined transport distances of lithic raw materials to understand patterns of movement, assessed the cultural capacities required to manufacture and use different sets of tools, applied stochastic models to examine the geographic distribution of sites, and reconstructed biome classes and climatic constraints. Our large-scale analysis allowed the research team to integrate different types of information and examine diachronic trends during the MSA. Based on our results, the range of cultural capacity expanded during the MSA. We define cultural capacity as the behavioral potential of a group expressed through the problem-solution distance required to manufacture and use tools. Our dataset also indicates that the actual behavior exhibited by MSA people, their cultural performance as expressed in the archaeological record, is not equivalent to their cultural capacity. Instead we observe that the main signature of the southern African MSA is its overall variability, as demonstrated by changing sets of cultural performances. Finally, at the scale of resolution considered here, our results suggest that climate is not the most significant factor driving human activities during the MSA. Instead, we postulate that behavioral flexibility itself became the key adaptation.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Institutional subscriptions

Fig. 1
Fig. 2
Fig. 3
Fig. 4
Fig. 5
Fig. 6
Fig. 7
Fig. 8
Fig. 9
Fig. 10

Similar content being viewed by others

References

  • Ambrose, S. H. (2006). Howiesons Poort lithic raw material procurement patterns and the evolution of modern human behavior: a response to Minichillo. Journal of Human Evolution, 50, 365–369.

    Article  Google Scholar 

  • Avery, D. M. (1982). The micromammalian fauna from border cave, Kwazulu, South Africa. Journal of Archaeological Science, 9, 187–204.

    Article  Google Scholar 

  • Avery, D. M. (1992). The environment of early modern humans at border cave, South Africa: micromammalian evidence. Palaeogeography, Palaeoclimatology, Palaeoecology, 91, 71–87.

    Article  Google Scholar 

  • Avery, G., Halkett, D., Orton, J., Steele, T., Tusenius, M., & Klein, R. G. (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 

  • Baxter, M. J. (2001). Methodological issues in the study of assemblage diversity. American Antiquity, 66, 715–725.

    Article  Google Scholar 

  • Beaumont, P. B. (1990). Kathu Pan. In P. Beaumont, & D. Morris (Eds.), Guide to archaeological sites in the Northern Cape (pp. 75–100). Kimberley: McGregor Museum.

  • Beven, K. J., & Kirkby, M. J. (1979). A physically-based variable contributing area model of basin hydrology. Hydrological Science Bulletin, 24, 43–69.

    Article  Google Scholar 

  • Binford, L. R. (1979). Organization and formation processes: looking at curated technologies. Journal of Anthropological Research, 35, 255–273.

    Article  Google Scholar 

  • Binford, L. R. (1980). Willow smoke and dogs’ tails: hunter-gatherer settlement systems and archaeological site formation. American Antiquity, 45, 4–20.

    Article  Google Scholar 

  • Binford, L. R. (1982). The archaeology of space. Journal of Anthropological Archaeology, 1, 5–31.

    Article  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 

  • Böhner, U. (2008). Die schicht E3 der sesselfelsgrotte und die funde aus dem abri I am schulerloch. Späte micoquien-inventare und ihre stellung zum moustérien. Stuttgart: Steiner.

    Google Scholar 

  • Böhner, J., & Antonić, O. (2009). Land surface parameters specific to topo-climatology. In T. Hengl & H. I. Reuter (Eds.), Geomorphometry: concepts, software, applications (pp. 195–226). Amsterdam: Elsevier.

    Chapter  Google Scholar 

  • Brew, J. O. (1946). The use and abuse of taxonomy: the archaeology of Alkali Ridge, Utah. Papers of the Peabody Museum of Archaeology and Ethnology, 21, 44–66.

    Google Scholar 

  • Brown, K. S., Marean, C. W., Herries, A. I. R., Jacobs, Z., Tribolo, C., Braun, D., Roberts, D. L., Meyer, C. M., & Bernatchez, J. (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., Bernatchez, J., Karkanas, P., & Matthews, T. (2012). An early and enduring advanced technology originating 71,000 years ago in South Africa. Nature, 491, 590–593.

    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 

  • Caley, T., Kim, J.-H., Malaizé, B., Giraudeau, J., Laepple, T., Caillon, N., Charlier, K., Rebaubier, H., Rossignol, L., Castañeda, I. S., Schouten, S., & Sinninghe Damsté, J. S. (2011). High-latitude obliquity as a dominant forcing in the Agulhas current system. Climate of the Past, 7, 1285–1296.

    Article  Google Scholar 

  • Carter, P. L., & Vogel, J. C. (1974). The dating of industrial assemblages from stratified sites in eastern Lesotho. Man, 9(4), 557–570.

    Article  Google Scholar 

  • Charrié-Duhaut, A., Porraz, G., Cartwright, C. R., Igreja, M., Connan, J., Poggenpoel, C., & Texier, P. J. (2013). First molecular identification of a hafting adhesive in the late Howiesons Poort at Diepkloof Rock Shelter (Western Cape, South Africa). Journal of Archaeological Science, 40, 3506–3518.

    Article  Google Scholar 

  • Chase, B. M. (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 dynamics of southern Africa’s winter rainfall zone. Earth-Science Reviews, 84, 103–138.

    Article  Google Scholar 

  • Clark, J. L., & Kandel, A. W. (2013). The evolutionary implications of variation in human hunting strategies and diet breadth during the Middle Stone Age of southern Africa. Current Anthropology, 54(8), 10.1086/673386.

  • Conard, N. J., Porraz, G., & Wadley, L. (2012). What is in a name? Characterising the post-Howieson’s Poort at Sibudu. South African Archaeological Bulletin, 67, 180–199.

    Google Scholar 

  • Conrad, O. (2007). SAGA–Entwurf, Funktionsumfang und Anwendung eines Systems für Automatisierte Geowissenschaftliche Analysen. Ph.D. dissertation, Göttingen: University of Göttingen.

  • d’Errico, F., Backwell, L., Villa, P., Degano, I., Lucejko, J. J., Bamford, M. K., Higham, T. F. G., Colombini, M. P., & Beaumont, P. B. (2012a). Early evidence of San material culture represented by organic artifacts from Border Cave, South Africa. Proceedings of the National Academy of Sciences of the United States of America, 109, 13214–13219.

  • d’Errico, F., Backwell, L. R., & Wadley, L. (2012b). Identifying regional variability in Middle Stone Age bone technology: the case of Sibudu cave. Journal of Archaeological Science, 39, 2479–2495.

    Article  Google Scholar 

  • Deacon, J. (1978). Changing patterns in the late Pleistocene/early Holocene prehistory of southern Africa as seen from the Nelson Bay Cave stone artifact sequence. Quaternary Research, 10, 84–111.

    Article  Google Scholar 

  • Deacon, H. J. (1979). Excavations at Boomplaas Cave—a sequence through the upper Pleistocene and Holocene in South Africa. World Archaeology, 10, 241–257.

    Article  Google Scholar 

  • Deacon, H. J. (1989). Late Pleistocene palaeoecology and archaeology in the Southern Cape, South Africa. In C. Stringer & P. Mellars (Eds.), The human revolution: behavioral and biological perspectives on the origins of modern humans (pp. 547–564). Princeton: Princeton University Press.

    Google Scholar 

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

    Article  Google Scholar 

  • d'Errico, F., & Henshilwood, C. (2007). Additional evidence for bone technology in the southern African Middle Stone Age. Journal of Human Evolution, 52, 142–163.

    Article  Google Scholar 

  • d'Errico, F., Henshilwood, C., & Nilssen, P. (2001). An engraved bone fragment from c. 70,000-year-old Middle Stone Age levels at Blombos Cave, South Africa: Implications for the origin of symbolism and language. Antiquity, 75, 309–318.

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

    Article  Google Scholar 

  • Dietl, H., Kandel, A. W., & Conard, N. J. (2005). Middle Stone Age settlement and land use at the open-air sites of Geelbek and Anyskop, South Africa. Journal of African Archaeology, 3(2), 233–244.

    Article  Google Scholar 

  • Elith, J., Leathwick, J. R., & Hastie, T. (2008). A working guide to boosted regression trees. Journal of Animal Ecology, 77, 802–813.

    Article  Google Scholar 

  • Evans, U. (1994). Hollow Rock Shelter, a Middle Stone Age site in the Cederberg. Southern African Field Archaeology, 3, 63–73.

    Google Scholar 

  • Fielding, A. H., & Bell, J. F. (1997). A review of methods for the assessment of prediction errors in conservation presence/absence models. Environmental Conservation, 24(1), 38–49.

    Article  Google Scholar 

  • Friedman, J. H. (1999). Stochastic gradient boosting. Technical Report, Department of Statistics, Stanford: Stanford University. http://www.salford-systems.com/treenet.html. Accessed 3 March 2015

  • Friedman, J. H. (2001). Greedy function approximation: a gradient boosting machine. Annals of Statistics, 29, 1189–1232.

    Article  Google Scholar 

  • Gelhausen, F. (2011). Siedlungsmuster der allerødzeitlichen Federmessergruppen in Niederbieber, Stadt Neuwied. Mainz: Verlag des Römisch-Germanischen Zentralmuseums.

  • Gibson, N. E., Wadley, L., & Williamson, B. S. (2004). Microscopic residues as evidence of hafting on backed tools from the 60 000 to 68 000 year-old Howiesons Poort layers of Rose Cottage Cave, South Africa. Southern African Humanities, 16, 1–11.

    Google Scholar 

  • Goldberg, P., Miller, C. E., Schiegl, S., Ligouis, B., Berna, F., Conard, N. J., & Wadley, L. (2009). Bedding, hearths, and site maintenance in the Middle Stone Age of Sibudu Cave, KwaZulu-Natal, South Africa. Archaeological and Anthropological Sciences, 1, 95–122.

    Article  Google Scholar 

  • Goodwin, A. J. H. (1929). The Stone Age cultures of South Africa. Annals of the South African Museum, 27, 95–145.

    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 

  • Haidle, M. N. (2009). How to think a simple spear? In S. A. de Beaune, F. L. Coolidge, & T. Wynn (Eds.), Cognitive archaeology and human evolution (pp. 57–73). New York: Cambridge University Press.

    Google Scholar 

  • Haidle, M. N. (2010). Working memory capacity and the evolution of modern cognitive capacities—implications from animal and early human tool use. Working memory: beyond language and symbolism Wenner-Gren Symposium. Current Anthropology, 51(S1), S149–S166.

    Article  Google Scholar 

  • Haidle, M. N. (2012). How to think tools? A comparison of cognitive aspects in tool behavior of animals and during human evolution. Cognitive perspectives in tool behavior, Vol. 1. http://tobias-lib.uni-tuebingen.de/frontdoor.php?source_opus=6014. Accessed 3 March 2015

  • Haidle, M. N., Bolus, M., Collard, M., Conard, N. J., Garofoli, D., Lombard, M., Nowell, A., Tennie, C., & Whiten, A. (2015). The nature of culture: an eight-grade model for the evolution and expansion of cultural capacities in hominins and other animals. Journal of Anthropological Sciences 93:1-28. doi:10.4436/jass.93011.

  • Haidle, M. N., & Conard, N. J. (2011). The nature of culture—summary report on an interdisciplinary symposium held in Tübingen, Germany, 15–18 June 2011. Mitteilungen der Gesellschaft für Urgeschichte, 20, 65–78.

    Google Scholar 

  • Haidle, M. N., Bolus, M., Bruch, A., Hertler, C., Kandel, A. W., Märker, M., Conard, N. J., Hochschild, V., Schrenk, F., & Mosbrugger, V. (2010). The role of culture in early expansions of humans—a new research center. Quaternary International, 223–224, 429–430.

    Article  Google Scholar 

  • Hanley, J. A., & McNeil, B. J. (1982). The meaning and use of the area under a receiver operating characteristic (ROC) curve. Radiology, 143, 29–36.

    Article  Google Scholar 

  • Henshilwood, C. S., & Dubreuil, B. (2011). The Still Bay and Howiesons Poort, 77–59 ka. Symbolic material culture and the evolution of the mind during the African Middle Stone Age. Current Anthropology, 52, 361–400.

    Article  Google Scholar 

  • Henshilwood, C. S., & Marean, C. W. (2006). Remodelling the origins of modern human behaviour. In H. Soodyall (Ed.), The human genome and Africa. Part 1: history and archaeology (pp. 31–48). Pretoria: Human Sciences Research Council.

    Google Scholar 

  • Henshilwood, C. S., Sealy, J. C., Yates, R., Cruz-Uribe, K., Goldberg, P., Grine, F. E., Klein, R. G., Poggenpoel, C., van Niekerk, K., & Watts, I. (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., Mercier, N., Sealy, J. C., Valladas, H., Watts, I., & Wintle, A. G. (2002). Emergence of modern human behavior: 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. L., & Jacobs, Z. (2004). Middle Stone Age shell beads from South Africa. Nature, 304, 404.

    Google Scholar 

  • Henshilwood, C. S., d'Errico, F., & Watts, I. (2009). Engraved ochers 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., Menu, M., & García Moreno, R. (2011). A 100,000-year-old ocher-processing workshop at Blombos Cave, South Africa. Science, 334, 219–222.

    Article  Google Scholar 

  • Henshilwood, C. H., van Niekerk, K. L., Wurz, S., Delagnes, A., Armitage, S. J., Rifkin, R. F., Douze, K., Keene, P., Haaland, M. M., Reynard, J., Discamps, E., & Mienies, S. S. (2014). Klipdrift Shelter, southern Cape, South Africa: preliminary report on the Howiesons Poort layers. Journal of Archaeological Science, 45, 284–303.

    Article  Google Scholar 

  • Hodgskiss, T. (2012). An investigation into the properties of the ochre from Sibudu, KwaZulu-Natal, South Africa. Southern African Humanities, 24, 99–120.

    Google Scholar 

  • Högberg, A., & Larsson, L. (2011). Lithic technology and behavioural modernity: new results from the Still Bay site, Hollow Rock Shelter, Western Cape Province, South Africa. Journal of Human Evolution, 61, 133–155.

    Article  Google Scholar 

  • Holzkämper, S., Holmgren, K., Lee-Thorp, J., Talma, S., Mangini, A., & Partridge, T. (2009). Late Pleistocene stalagmite growth in Wolkberg cave, South Africa. Earth and Planetary Science Letters, 282, 212–221.

    Article  Google Scholar 

  • Hosmer, D. W., & Lemeshow, S. (2000). Applied logistic regression (2nd ed.). New York: Wiley.

    Book  Google Scholar 

  • Jacobs, Z., Roberts, R. G., Galbraith, R. F., Deacon, H. J., Grün, R., Mackay, A., Mitchell, P., Vogelsang, R., & Wadley, L. (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, South Africa. Journal of Archaeological Science, 35, 1790–1807.

    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 

  • Jarvis, A., Reuter, H. I., Nelson, A., & Guevara, E. (2006). Hole-filled seamless SRTM data V3, International Centre for Tropical Agriculture (CIAT), http://srtm.csi.cgiar.org. Accessed 3 March 2015

  • 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 

  • Kandel, A. W., & Conard, N. J. (2012). Settlement patterns during the Earlier and Middle Stone Age around Langebaan Lagoon, Western Cape (South Africa). Quaternary International, 270, 15–29. doi:10.1016/j.quaint.2011.06.038.

  • Kaplan, J. (1990). The Umhlatuzana Rock Shelter sequence: 100,000 years of Stone Age history. Natal Museum Journal of Humanities, 2, 1–94.

    Google Scholar 

  • Kaufman, D. (1998). Measuring archaeological diversity: an application of the jackknife technique. American Antiquity, 63, 73–85.

    Article  Google Scholar 

  • Kintigh, K. W. (1984). Measuring archaeological diversity by comparison with simulated assemblages. American Antiquity, 49, 44–54.

    Article  Google Scholar 

  • Klein, R. G. (1974). Environment and subsistence of prehistoric man in the Southern Cape Province, South Africa. World Archaeology, 5, 249–284.

    Article  Google Scholar 

  • Klein, R. G. (1977). The mammalian fauna from the Middle and Later Stone Age (Later Pleistocene) levels of Border Cave, Natal Province, South Africa. South African Archaeological Bulletin, 32, 14–27.

    Article  Google Scholar 

  • Klein, R. G. (2001). Southern Africa and modern human origins. Journal of Anthropological Research, 57, 1–16.

    Article  Google Scholar 

  • Klein, R. G. (2008). Out of Africa and the evolution of human behavior. Evolutionary Anthropology, 17, 267–281.

    Article  Google Scholar 

  • Klein, R. G., Avery, G., Cruz-Uribe, K., & Steele, T. E. (2007). The mammalian fauna associated with an archaic hominin skullcap and later Acheulean artifacts at Elandsfontein, Western Cape Province, South Africa. Journal of Human Evolution, 52, 164–186.

    Article  Google Scholar 

  • Kovarovic, K., Aiello, L. C., Cardini, A., & Lockwood, C. A. (2011). Discriminant function analyses in archaeology: are classification rates too good to be true? Journal of Archaeological Science, 38, 3006–3018.

    Article  Google Scholar 

  • Kuman, K., & Clarke, R. J. (2000). Stratigraphy, artefact industries and hominid associations for Sterkfontein, Member 5. Journal of Human Evolution, 38, 827–847.

    Article  Google Scholar 

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

    Article  Google Scholar 

  • Langejans, G. H. J., van Niekerk, K. L., Dusseldorp, G. L., & Thackeray, J. F. (2012). Middle Stone Age shellfish exploitation: Potential indications for mass collecting and resource intensification at Blombos Cave and Klasies River, South Africa. Quaternary International, 270, 80–94.

    Article  Google Scholar 

  • Lombard, M. (2006). First impressions of the functions and hafting technology of Still Bay pointed artefacts from Sibudu Cave. Southern African Humanities, 18(1), 27–41.

    Google Scholar 

  • Lombard, M. (2008). Finding resolution for the Howiesons Poort through the microscope: micro-residue analysis of segments from Sibudu Cave, South Africa. Journal of Archaeological Science, 35, 26–41.

    Article  Google Scholar 

  • Lombard, M. (2011). Quartz-tipped arrows older than 60 ka: further use-trace evidence from Sibudu, KwaZulu-Natal, South Africa. Journal of Archaeological Science, 38, 1918–1930.

    Article  Google Scholar 

  • Lombard, M. (2012). Thinking through the Middle Stone Age of sub-Saharan Africa. Quaternary International, 270, 140–155.

    Article  Google Scholar 

  • Lombard, M., & Haidle, M. N. (2012). Thinking a bow-and-arrow: cognitive implications of Middle Stone Age bow and stone-tipped arrow technology. Cambridge Archaeological Journal, 22, 237–264.

    Article  Google Scholar 

  • Lombard, M., & Parsons, I. (2011). What happened to the human mind after the Howiesons Poort? Antiquity, 85, 1433–1443.

    Article  Google Scholar 

  • Lombard, M., & Phillipson, L. (2010). Indications of bow and stone-tipped arrow use 64000 years ago in KwaZulu-Natal, South Africa. Antiquity, 84, 1–14.

    Article  Google Scholar 

  • Lombard, M., Wadley, L., Jacobs, Z., Mohapi, M., & Roberts, R. G. (2010). Still bay and serrated points from Umhlatuzana Rock Shelter, Kwazulu-Natal, South Africa. Journal of Archaeological Science, 37, 1–12.

    Article  Google Scholar 

  • Lombard, M., Wadley, L., Deacon, J., Wurz, S., Parsons, I., Mohapi, M., Swart, J., & Mitchell, P. (2012). South African and Lesotho Stone Age sequence updated (I). South African Archaeological Bulletin, 67, 123–144.

    Google Scholar 

  • Mackay, A. (2009). History and selection in the late Pleistocene archaeology of the Western Cape, South Africa. Unpublished Ph.D. thesis, Canberra: Australian National University.

  • Mackay, A. (2011a). Potentially stylistic differences between backed artefacts from two nearby sites occupied ~60,000 years before present in South Africa. Journal of Anthropological Archaeology, 30, 235–245.

    Article  Google Scholar 

  • Mackay, A. (2011b). Nature and significance of the Howiesons Poort to post-Howiesons Poort transition at Klein Kliphuis rockshelter, South Africa. Journal of Archaeological Science, 38, 1430–1440.

    Article  Google Scholar 

  • Mackay, A., & Welz, A. (2008). Engraved ocher from a Middle Stone Age context at Klein Kliphuis in the Western Cape of South Africa. Journal of Archaeological Science, 35, 152–1532.

    Google Scholar 

  • Mackay, A. Stewart, B. A., & Chase, B. M. (2014). Coalescence and fragmentation in the late Pleistocene archaeology of southernmost Africa. Journal of Human Evolution, 10.1016/j.jhevol.2014.03.003.

  • Malan, B. D. (1955). The archaeology of the Tunnel Cave and Skildergat Kop, Fish Hoek, Cape of Good Hope. South African Archaeological Bulletin, 10, 3–9.

    Article  Google Scholar 

  • Manel, S., Williams, H. C., & Ormerod, S. J. (2001). Evaluating presence–absence models in ecology: the need to account for prevalence. Journal of Applied Ecology, 38, 921–931.

    Article  Google Scholar 

  • Marean, C. W., & Assefa, Z. (2005). The Middle and Upper Pleistocene African record for the biological and behavioral origins of modern humans. In A. B. Stahl (Ed.), African archaeology (pp. 93–129). Malden: Blackwell.

    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, 1, 14–83.

    Google Scholar 

  • Marean, C. W., Bar-Matthews, M., Bernatchez, J., Fisher, E., Goldberg, P., Herries, A. I. R., Jacobs, Z., Jerardino, A., Karkanas, P., Minichillo, T., Nilssen, P. J., Thompson, E., Watts, I., & Williams, H. M. (2007). Early human use of marine resources and pigment in South Africa during the Middle Pleistocene. Nature, 449, 905–908.

    Article  Google Scholar 

  • Marean, C. W., Bar-Matthews, M., Fisher, E., Goldberg, P., Herries, A., Karkanas, P., Nilssen, P. J., & Thompson, E. (2010). The stratigraphy of the Middle Stone Age sediments at Pinnacle Point Cave 13B (Mossel Bay, Western Cape Province, South Africa). Journal of Human Evolution, 59, 234–255.

    Article  Google Scholar 

  • Märker, M., Bolus, M., & Kanaeva, Z. (2011). Spatial assessment of early human expansions using GIS and Database Techniques: Examples from Southern Africa and Eurasia. In R. Macchiarelli, & G.-C. Weniger (Eds.), Pleistocene databases: Acquisition, storing, sharing (pp. 69–75). Wissenschaftliche Schriften des Neanderthal Museums 4. Mettmann: Neanderthal Museum.

  • Mason, R. J. (1962). Prehistory of the Transvaal. Johannesburg: Witwatersrand University Press.

    Google Scholar 

  • Mazza, P., Martini, F., Sala, B., Magi, M., Colombini, M. P., Giachi, G., Landucci, F., Lemorini, C., Modugno, F., & Ribechini, E. (2006). A new Palaeolithic discovery: tar-hafted stone tools in a European Mid-Pleistocene bone-bearing bed. Journal of Archaeological Science, 33, 1310–1318.

    Article  Google Scholar 

  • McBrearty, S. (2012). Sharpening the mind. Nature, 491, 531–532.

    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 

  • McCall, G. S. (2007). Behavioral ecological models of lithic technological change during the later Middle Stone Age of South Africa. Journal of Archaeological Science, 34, 1738–1751.

    Article  Google Scholar 

  • McCartney, P. H., & Glass, M. F. (1990). Simulation models and the interpretation of archaeological diversity. American Antiquity, 55, 521–536.

    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 

  • Mitchell, P. J. (1996). The late Quaternary of the Lesotho highlands, southern Africa: Preliminary results and future potential of ongoing research at Sehonghong shelter. Quaternary International, 33, 35–43.

    Article  Google Scholar 

  • Mitchell, P. J., & Steinberg, J. M. (1992). Ntloana Tsoana: A Middle Stone Age sequence from western Lesotho. South African Archaeological Bulletin, 47, 26–33.

    Article  Google Scholar 

  • Moore, I. D., Garson, R. B., & Ladson, A. R. (1991). Digital terrain modelling: a review of hydrological, geomorphological and biological applications. Hydrological Processes, 5, 3–30.

    Article  Google Scholar 

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

    Article  Google Scholar 

  • Nobre, A. D., Cuartas, L. A., Hodnett, M., Rennó, C. D., Rodrigues, G., Silveira, A., Waterloo, M., & Saleska, S. (2011). Height Above the Nearest Drainage—a hydrologically relevant new terrain model. Journal of Hydrology, 404(1–2), 13–29.

    Article  Google Scholar 

  • Olaya, V., & Conrad, O. (2008). Geomorphometry in SAGA. In T. Hengl & H. I. Reuter (Eds.), Geomorphometry: concepts, software, applications (pp. 293–308). Amsterdam: Elsevier.

    Google Scholar 

  • Olson, D. M., Dinerstein, E., Wikramanayake, E. D., Burgess, N. D., Powell, G. V. N., Underwood, E. C., D'amico, J. A., Itoua, I., Strand, H. E., Morrison, J. C., Loucks, C. J., Allnutt, T. F., Ricketts, T. H., Kura, Y., Lamoreux, J. F., Wettengel, W. W., Hedao, P., & Kassem, K. R. (2001). Terrestrial ecoregions of the world: a new map of life on Earth. BioScience, 51, 933–938.

    Article  Google Scholar 

  • Opperman, H., & Heydenrych, B. (1990). A 22 000 year-old Middle Stone Age camp site with plant food remains from the North-Eastern Cape. South African Archaeological Bulletin, 45, 93–99.

    Article  Google Scholar 

  • Parkington, J. E. (2003). Middens and moderns: shellfishing and the Middle Stone Age of the Western Cape, South Africa. South African Journal of Science, 99, 243–247.

    Google Scholar 

  • Planchon, O., & Darboux, F. (2001). A fast, simple and versatile algorithm to fill the depressions of digital elevation models. Catena, 46, 159–176.

    Article  Google Scholar 

  • Porraz, G., Texier, P.-J., Rigaud, J.-P., Parkington, J. E., Poggenpoel, C., & Roberts, D. L. (2008). Preliminary characterization of a Middle Stone Age lithic assemblage preceding the 'classic' Howieson's Poort complex at Diepkloof Rock Shelter, Western Cape Province, South Africa. South African Archaeological Society Goodwin Series, 10, 105–121.

    Google Scholar 

  • Porraz, G., Texier, P.-J., Archer, W., Piboule, M., Rigaud, J.-P., & Tribolo, C. (2013). Technological successions in the Middle Stone Age sequence of Diepkloof Rock Shelter, Western Cape, South Africa. Journal of Archaeological Science, 40, 3376–3400.

    Article  Google Scholar 

  • Rector, A. L., & Reed, K. E. (2010). Middle and late Pleistocene faunas of Pinnacle Point and their paleoecological implications. Journal of Human Evolution, 59, 340–357.

    Article  Google Scholar 

  • Reed, K. E. (1997). Early hominid evolution and ecological change through the African Plio-Pleistocene. Journal of Human Evolution, 32, 289–322.

    Article  Google Scholar 

  • Reed, K. E. (1998). Using large mammal communities to examine ecological and taxonomic structure and predict vegetation in extant and extinct assemblages. Paleobiology, 24, 384–408.

    Google Scholar 

  • Reed, K. E., & Rector, A. L. (2007). African Pliocene Paleoecology. Hominin habitats, resources, and diets. In P. S. Ungar (Ed.), Evolution of the Human Diet (pp. 262–288). New York: Oxford University Press.

    Google Scholar 

  • Reineking, B., & Schröder, B. (2006). Constrain to perform: regularization of habitat models. Ecological Modeling, 193, 675–690.

    Article  Google Scholar 

  • Rhode, D. (1988). Measurement of archaeological diversity and the sample-size effect. American Antiquity, 53, 708–716.

    Article  Google Scholar 

  • Richter, J. (1990). Diversität als Zeitmass im Spätmagdalénien. Archäologisches Korrespondenzblatt, 20, 249–257.

    Google Scholar 

  • Rigaud, J.-P., Texier, P.-J., Parkington, J. E., & Poggenpoel, C. (2006). Le mobilier Still Bay et Howiesons Poort de l'abri Diepkloof. La chronologie du Middle Stone Age sud-africain et ses implications. Comptes Rendus Palevol, 5, 839–849.

    Article  Google Scholar 

  • Rots, V., & Van Peer, P. (2006). Early evidence of complexity in lithic economy: core-axe production, hafting and use at Late Middle Pleistocene site 8-B-11, Sai Island (Sudan). Journal of Archaeological Science, 33, 360–371.

    Article  Google Scholar 

  • Sappington, J. M., Longshore, K. M., & Thompson, D. B. (2007). Quantifying landscape ruggedness for animal habitat analysis: a case study using bighorn sheep in the Mojave Desert. Journal of Wildlife Management, 71(5), 1419–1426.

    Article  Google Scholar 

  • Schmidt, P., Porraz, G., Slodczyk, A., Bellot-gurlet, L., Archer, W., & Miller, C. E. (2013). Heat treatment in the South African Middle Stone Age: temperature induced transformations of silcrete and their technological implications. Journal of Archaeological Science, 40, 3519–3531.

    Article  Google Scholar 

  • Sealy, J. C. (2006). Diet, mobility, and settlement pattern among Holocene hunter-gatherers in southernmost Africa. Current Anthropology, 47, 569–595.

    Article  Google Scholar 

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

    Google Scholar 

  • Soriano, S., Villa, P., & Wadley, L. (2007). Blade technology and tool forms in the Middle Stone Age of South Africa: the Howiesons Poort and post-Howiesons Poort at Rose Cottage Cave. Journal of Archaeological Science, 34, 681–703.

    Article  Google Scholar 

  • Soriano, S., Villa, P., & Wadley, L. (2009). Ocher for the toolmaker: shaping the Still Bay points at Sibudu (KwaZulu-Natal, South Africa). Journal of African Archaeology, 7, 41–54.

    Article  Google Scholar 

  • Steele, T. E., & Klein, R. G. (2005). Mollusk and tortoise size as proxies for stone age population density in South Africa: Implications for the evolution of human cultural capacity. Munibe. Sociedad de Ciencias Naturales Aranzadi (San Sebastian), 57, 221–237.

    Google Scholar 

  • Steele, T. E., & Klein, R. G. (2009). Late Pleistocene subsistence strategies and resource intensification in Africa. In J.-J. Hublin & M. P. Richards (Eds.), The evolution of hominin diets: integrating approaches to the study of Palaeolithic subsistence, vertebrate paleobiology and paleoanthropology (pp. 113–126). Dordrecht: Springer.

    Chapter  Google Scholar 

  • Steele, T. E., & Klein, R. G. (2013). The Middle and Later Stone Age faunal remains from Diepkloof Rockshelter, Western Cape, South Africa. Journal of Archaeological Science, 40, 3453–3462.

    Article  Google Scholar 

  • Steele, T. E., Mackay, A., Orton, J., & Schwortz, S. (2012). Varsche Rivier 003, a new Middle Stone Age site in southern Namaqualand, South Africa. South African Archaeological Bulletin, 67, 108–119.

    Google Scholar 

  • Stewart, B. A., Dewar, G. I., Morley, M. W., Inglis, R. H., Wheeler, M., Jacobs, Z., & Roberts, R. G. (2012). Afromontane foragers of the Late Pleistocene: site formation, chronology and occupational pulsing at Melikane Rockshelter, Lesotho. Quaternary International, 270, 40–60.

    Article  Google Scholar 

  • Swets, J. A. (1988). Measuring the accuracy of diagnostic systems. Science, 240, 1285–1293.

    Article  Google Scholar 

  • Tattersall, I. (2014). An evolutionary context for the emergence of language. Language Sciences, 46B, 199–206.

    Article  Google Scholar 

  • Texier, P.-J., Porraz, G., Parkington, J. E., Rigaud, J.-P., Poggenpoel, C., Miller, C. E., Tribolo, C., Cartwright, C., Coudenneau, A., Klein, R. G., Steele, T., & Verna, C. (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 Sciences of the United States of America, 107, 6180–6185.

    Article  Google Scholar 

  • Texier, P. J., Porraz, G., Parkington, J., Rigaud, J.-P., Poggenpoel, C., & Tribolo, C. (2013). The context, form and significance of the MSA engraved ostrich eggshell collection from Diepkloof Rock Shelter, Western Cape, South Africa. Journal of Archaeological Science, 40, 3412–3431.

    Article  Google Scholar 

  • Thackeray, J. F. (1988). Molluscan fauna from Klasies River, South Africa. South African Archaeological Bulletin, 43, 27–32.

    Article  Google Scholar 

  • Thackeray, A. I. (2000). Middle Stone Age artifacts from the 1993 and 1995 excavations of Die Kelders Cave 1, South Africa. Journal of Human Evolution, 38, 147–168.

    Article  Google Scholar 

  • Thieme, H. (1999). Altpaläolithische Holzgeräte aus Schöningen. Lkr. Helmstedt. Germania, 77(2), 451–487.

    Google Scholar 

  • Tribolo, C., Mercier, N., Valladas, H., Joron, J. L., Guibert, P., Lefrais, Y., Selo, M., Texier, P.-J., Rigaud, J.-P., Porraz, G., Poggenpoel, C., Parkington, J., Texier, J.-P., & Lenoble, A. (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., Cantin, N., Lefrais, Y., Miller, C. E., Porraz, G., Parkington, J., Rigaud, J.-P., & Texier, P.-J. (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 

  • Turner, A. (1999). Evolution in the Africa Plio-Pleistocene Mammalian fauna: correlation and causation. In T. G. Bromage & F. Schrenk (Eds.), African biogeography, climate change, and human evolution (pp. 76–90). New York: Oxford University Press.

    Google Scholar 

  • Underhill, D. (2011). A history of Stone Age archaeological study in South Africa. South African Archaeological Bulletin, 66, 3–14.

    Google Scholar 

  • Uthmeier, T. (2004). Micoquien, Aurignacien und Gravettien in Bayern. Eine regionale Studie zum Übergang vom Mittel- zum Jungpaläolithikum. Archäologische Berichte, 18. Bonn: Habelt.

    Google Scholar 

  • Vanhaeren, M., d’Errico, F., van Niekerk, K. L., Henshilwood, C. S., & Erasmus, R. M. (2013). Thinking strings: additional evidence for personal ornament use in the Middle Stone Age at Blombos Cave, South Africa. Journal of Human Evolution, 64, 500–517.

    Article  Google Scholar 

  • Villa, P., Delagnes, A., & Wadley, L. (2005). A late Middle Stone Age artifact assemblage from Sibudu (KwaZulu-Natal): comparisons with the European Middle Paleolithic. Journal of Archaeological Science, 32, 399–422.

    Article  Google Scholar 

  • Villa, P., Soressi, M., Henshilwood, C. S., & Mourre, V. (2009). The Still Bay points of Blombos Cave (South Africa). Journal of Archaeological Science, 36, 441–460.

    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 

  • Vogelsang, R. (1998). Middle-Stone-Age-Fundstellen in Südwest-Namibia. Köln: Heinrich-Barth-Institute.

    Google Scholar 

  • Vogelsang, R., Richter, J., Jacobs, Z., Eichhorn, B., Linseele, V., & Roberts, R. G. (2010). New excavations of Middle Stone Age deposits at Apollo 11 Rockshelter, Namibia: stratigraphy, archaeology, chronology and past environments. Journal of African Archaeology, 8(2), 185–218.

    Article  Google Scholar 

  • Volman, T. P. (1978). Early archeological evidence for shellfish collecting. Science, 201, 911–913.

    Article  Google Scholar 

  • Volman, T. P. (1981). The Middle Stone Age in the Southern Cape. Ph.D. dissertation, Chicago: University of Chicago.

  • Wadley, L. (1991). Rose Cottage Cave: background and a preliminary report on the recent excavations. South African Archaeological Bulletin, 46, 125–130.

    Article  Google Scholar 

  • Wadley, L. (2005). Putting ocher to the test: replication studies of adhesives that may have been used for hafting tools in the Middle Stone Age. Journal of Human Evolution, 49, 587–601.

    Article  Google Scholar 

  • Wadley, L. (2006). Partners in grime: results of multi-disciplinary archaeology at Sibudu Cave. Southern African Humanities, 18(1), 315–341.

    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. (2010a). Compound-adhesive manufacture as a behavioral proxy for complex cognition in the Middle Stone Age. Current Anthropology, 51(S1), S111–S119.

    Article  Google Scholar 

  • Wadley, L. (2010b). Were snares and traps used in the Middle Stone Age and does it matter? A review and a case study from Sibudu, South Africa. Journal of Human Evolution, 58, 179–192.

    Article  Google Scholar 

  • Wadley, L. (2013). Recognizing complex cognition through innovative technology in Stone Age and Palaeolithic sites. Cambridge Archaeological Journal, 23, 163–183.

    Article  Google Scholar 

  • Wadley, L., & Mohapi, M. (2008). A segment is not a monolith: evidence from the Howiesons Poort of Sibudu, South Africa. Journal of Archaeological Science, 35, 2594–2605.

    Article  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 

  • Wendt, W. E. (1976). ‘Art mobilier’ from the Apollo 11 Cave, South West Africa: Africa's oldest dated works of art. South African Archaeological Bulletin, 31, 5–11.

    Article  Google Scholar 

  • Weniger, G. C. (1991). Überlegungen zur Mobilität jägerischer Gruppen im Jungpaläolithikum. In Urgeschichte als Kulturanthropologie. Beiträge zum 70. Geburtstag von Karl J. Narr. Saeculum, 42, 82–103.

    Article  Google Scholar 

  • Wilkins, J., Schoville, B. J., Brown, K. S., & Chazan, M. (2012). Evidence for early hafted hunting technology. Science, 338, 942–946.

    Article  Google Scholar 

  • Will, M., Parkington, J. E., Kandel, A. W., & Conard, N. J. (2013). Coastal adaptations and the MSA lithic assemblages from Hoedjiespunt 1 in the Western Cape, South Africa. Journal of Human Evolution, 64, 518–537.

    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., Le Roux, N. J., Gardner, S., & Deacon, H. J. (2003). Discriminating between the end products of the earlier Middle Stone Age sub-stages at Klasies River using biplot methodology. Journal of African Archaeology, 30, 1107–1126.

    Google Scholar 

  • Yokoyama, R., Shirasawa, M., & Pike, R. J. (2002). Visualizing topography by openness: a new application of image processing to digital elevation models. Photogrammetric Engineering & Remote Sensing, 68, 257–266.

    Google Scholar 

  • Zevenbergen, L. W., & Thorne, C. R. (1987). Quantitative analysis of the land surface topography. Earth Surface Process and Landform, 12, 47–56.

    Article  Google Scholar 

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

Download references

Acknowledgments

We thank the following staff and students at the University of Tübingen and the Senckenberg Research Institute in Frankfurt am Main: Krista Dotzel, Michaela Ecker, Matthias Göden, Maria Malina, Patrick Schmidt, Andreas Taller, and Manuel Will for data entry and quality control; Zara Kanaeva for preparation of database queries; Geraldine Quénéhervé for geographical analysis and map generation; Regine Stolarczyk for computational assistance with regards to tool specialization and tool groups; and Rebekka Volmer for assistance with the faunal data. We are grateful to Nicholas Conard, Manuel Will, and three anonymous reviewers for their constructive comments, criticisms, and complaints which helped improve this manuscript. In the end, of course, we take full responsibility for its content. The ROCEEH research project is funded by the Heidelberg Academy of Sciences and Humanities with support from the Senckenberg Research Institute and the University of Tübingen. Finally, we thank Iain Davidson, Leslie Aiello, and Dominique Grimaud-Hervé, who encouraged us to take chances we normally would not.

Author information

Authors and Affiliations

Authors

Corresponding author

Correspondence to Andrew W. Kandel.

Electronic Supplementary Material

Below is the link to the electronic supplementary material.

Table ESM 1

List of the localities and assemblages considered in the calculation of the Specialization Index. For Early and Late MSA, square brackets indicate the cultural designation as cited in the literature. (DOCX 18 kb)

Table ESM 2

List of the localities and assemblages considered in the calculation of the Tool Group Index. For Early and Late MSA, square brackets indicate the cultural designation as cited in the literature. (DOCX 20 kb)

Table ESM 3

List of the localities and assemblages considered in the calculation of rock transport distance. For Early and Late MSA, square brackets indicate the cultural designation as cited in the literature. (DOCX 18 kb)

Table ESM 4

Species-specific ecological profiles. Diet classes: HFR=frugivore; HXB=folivore/browser; HMB=mixed feeder; HMG=fresh grass grazer; HXG=obligate grazer. Body mass classes [kg]: 2b=2-5; 2c=5-10; 3a=10-20; 3b=20-50; 3c=50-100; 4a=100-200; 4b=200-500; 4c=500-1000; 5a=1000-2000; 5b=2000-5000. Locomotion types: ARB=arboreal; SAR=semi-arboreal/scansorial; AMP=amphibian; UBI=ubiquitous terrestrial; UNG=ungulate; GRA=graviportal. (DOCX 18 kb)

Table ESM 5

Characteristics of the six specialized herbivore communities in the reference sample based on data derived from 30 national parks in southern Africa. The classification results from faunal datasets: d eco =ecological diversities; d rel =relative diversities, proxy for structural redundancy. Description of the sample: HFR=frugivore; HXB=folivore/browser; HMB=mixed feeder; HMG=fresh grass grazer; HXG=obligate grazer; SM= small <200 kg body mass; LA= large ≥200 kg. (DOCX 19 kb)

Table ESM 6

List of the localities and assemblages considered in assessment of fauna. For Early and Late MSA, square brackets indicate the cultural designation as cited in the literature. (DOCX 16 kb)

Table ESM 7

Quantification of specialized herbivore communities with regard to climatic regimes and vegetation characteristics. The classification results from faunal datasets: d eco =ecological diversities; d rel =relative diversities, proxy for structural redundancy. Description of the sample: HFR=frugivore; HXB=folivore/browser; HMB=mixed feeder; HMG=fresh grass grazer; HXG=obligate grazer; SM= small <200 kg body mass; LA= large ≥200 kg. Climatic regime: MAT=mean annual temperature; ∆T (WAQ-CQ)=temperature difference between mean temperature of the warmest and coldest quarter, proxy for temperature seasonality; MAP=mean annual precipitation; ∆P1 (WEQ-DQ)=precipitation difference between mean precipitation of wettest and driest quarter, proxy for precipitation seasonality; ∆P2 (WAQ-CQ)=precipitation difference between mean precipitation of warmest and coldest quarter, proxy for summer (>0) vs. winter rains (<0). Vegetation characteristics: LAI=leaf area index (leaf area in relation to base area); greenness=proxy for photosynthesis activity (varies between 0 and 1); vegetation cover=proxy for vegetation density (varies between 0 and 1). (DOCX 18 kb)

Table ESM 8

Classification functions for both sets of variables. See Table ESM-7 for legend. (DOCX 19 kb)

ESM 9

(DOCX 38 kb)

Rights and permissions

Reprints and permissions

About this article

Check for updates. Verify currency and authenticity via CrossMark

Cite this article

Kandel, A.W., Bolus, M., Bretzke, K. et al. Increasing Behavioral Flexibility? An Integrative Macro-Scale Approach to Understanding the Middle Stone Age of Southern Africa. J Archaeol Method Theory 23, 623–668 (2016). https://doi.org/10.1007/s10816-015-9254-y

Download citation

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s10816-015-9254-y

Keywords

Navigation