Microwear textures of Australopithecus africanus and Paranthropus robustus molars in relation to paleoenvironment and diet (original) (raw)

Abstract

The importance of diet in primate ecology has motivated the use of a variety of methods to reconstruct dietary habits of extinct hominin taxa. Dental microwear is one such approach that preserves evidence from consumed food items. This study is based on 44 specimens of Australopithecus africanus from Makapansgat and Sterkfontein, and 66 specimens of Paranthropus robustus from Swartkrans, Kromdraai and Drimolen. These samples enable examination of potential differences between the two assemblages of A. africanus, and among the various assemblages of P. robustus in relation to the paleoenvironmental reconstructions that have been proffered for each fossil site. Sixteen microwear texture variables were recorded for each specimen from digital elevation models generated using a white-light confocal profiler. Only two of these differ significantly between the Makapansgat and Sterkfontein samples of A. africanus. None of the microwear texture variables differs significantly among the samples of P. robustus. On the other hand, P. robustus has significantly higher values than A. africanus for 11 variables related to feature complexity, size, and depth; P. robustus exhibits rougher surfaces that comprise larger, deeper features. In contrast, A. africanus has smoother, simpler wear surfaces with smaller, shallower and more anisotropic features. As for possible habitat differences among the various sites, only a relatively small number of subtle differences are evident between the specimens of A. africanus from Makapansgat and Sterkfontein, and there are none among the specimens of P. robustus from various deposits. As such, it is reasonable to conclude that, while subtle differences in microwear textures may reflect differences in background habitats, the wear fabric differences between P. robustus and A. africanus are most reasonably interpreted as having been driven by dietary differences.

Loading...

Loading Preview

Sorry, preview is currently unavailable. You can download the paper by clicking the button above.

References (216)

  1. Adams, J.W., Rovinsky, D.S., Herries, A.I.R., Menter, C.G., 2016. Macromammalian faunas, biochronology and palaeoecology of the early Pleistocene Main Quarry hominin-bearing deposits of the Drimolen palaeocave system, South Africa. PeerJ 4, e1941.
  2. Arman, S.D., Ungar, P.S., Brown, C.A., DeSantis, L.R.G., Schmidt, C., Prideaux, G.J., 2016. Minimizing inter-microscope variability in dental microwear texture analysis. Surface Topography: Metrology and Properties 4, 024007.
  3. Avery, D.M., 1995. Southern savannas and Pleistocene hominid adaptations: the micromammalian perspective. In: Vrba, E.S., Denton, G.H., Partridge, T.C., Burckle, L.H. (Eds.), Paleoclimate and Evolution with Special Emphasis on Hu- man Origins. Yale University Press, New Haven, pp. 459e477.
  4. Avery, D.M., 1998. An assessment of the lower Pleistocene micromammalian fauna from Swartkrans Members 1e3, Gauteng, South Africa. Geobios 31, 393e414.
  5. Avery, D.M., 2001. The Plio-Pleistocene vegetation and climate of Sterkfontein and Swartkrans, South Africa, based on micromammals. Journal of Human Evolution 41, 113e132.
  6. Avery, D.M., 2007. Micromammals as palaeoenvironmental indicators of the southern African Quaternary. Transactions of the Royal Society of South Africa 62, 17e23.
  7. Avery, D.M., Stratford, D.J., S en egas, F., 2010. Micromammals and the formation of the Name Chamber, at Sterkfontein, South Africa. Geobios 43, 379e387.
  8. Bailey, G.N., Reynolds, S.C., King, G.C.P., 2011. Landscapes of human evolution: models and methods of tectonic geomorphology and the reconstruction of hominin landscapes. Journal of Human Evolution 60, 257e280.
  9. Bamford, M., 1999. Pliocene fossil woods from an early hominid cave deposit, Sterkfontein, South Africa. South African Journal of Science 95, 231e237.
  10. Bamford, M., 2000. Palaeoenvironmental reconstruction of the Upper Pliocene Sterkfontein site from fossil wood. In: Abstracts from the XVth INQUA Congress Held in Durban, Sosuth Africa, 3e11 August 1999. Quarternary International 63/ 64, 16.
  11. Behrensmeyer, A.K., 1982. Time resolution in fluvial vertebrate assemblages. Paleobiology 8, 211e227.
  12. Behrensmeyer, A.K., Bobe, R., Alemseged, Z., 2007. Approaches to the analysis of faunal change during the East African Pliocene. In: Bobe, R., Alemseged, Z., Behrensmeyer, A.K. (Eds.), Hominin Environments in the East African Pliocene: An Assessment of the Faunal Evidence. Springer, Dordrecht, pp. 1e24.
  13. Benefit, B.R., McCrossin, M.L., 1990. Diet, species diversity, and distribution of Af- rican fossil baboons. Kroeber Anthropological Society Papers 71e72, 79e93.
  14. Birchette, M., 1981. Postcranial remains of Cercopithecoides. American Journal of Physical Anthropology 54, 201.
  15. Birchette, M., 1982. The postcranial skeleton of Paracolobus chemeroni. Ph.D. Dissertation, Harvard University.
  16. Brain, C.K., 1958. The Transvaal ape-man-bearing cave deposits. Ph.D. Dissertation, University of Cape Town.
  17. Brain, C.K., 1975. An interpretation of the bone assemblage from the Kromdraai australopithecine site, South Africa. In: Tuttle, R.H. (Ed.), Paleanthropology, Morphology and Palaeoecology. Mouton, The Hague, pp. 225e243.
  18. Brain, C.K., 1976. A re-interpretation of the Swartkrans site and its remains. South African Journal of Science 72, 141e146.
  19. Brain, C.K., 1981. The Hunters or the Hunted? An Introduction to African Cave Taphonomy. University of Chicago Press, Chicago.
  20. Brain, C.K., 1988. New information from the Swartkrans Cave of relevance to "robust" australopithecines. In: Grine, F.E. (Ed.), Evolutionary History of the "Robust" Australopithecines. Aldine de Gruyter, New York, pp. 311e316.
  21. Brain, C.K., 1993. Structure and stratigraphy of the Swartkrans Cave in light of the new excavations. In: Brain, C.K. (Ed.), Swartkrans: A Cave's Chronicle of Early Man, Transvaal Museum Monograph No. 8. Transvaal Museum, Pretoria, pp. 23e34.
  22. Brain, C.K., 1994. The Swartkrans Paleontological Research Project in perspective e results and conclusions. South African Journal of Science 90, 220e223.
  23. Brain, C.K., Watson, V., 1992. A guide to the Swartkrans early hominid cave site. Annals of the Transvaal Museum 35, 343e365.
  24. Brain, C.K., Churcher, C.S., Clark, J.D., Grine, F.E., Shipman, P., Susman, R.L., Turner, A., Watson, V., 1988. New evidence of early hominids, their culture and environ- ment from the Swartkrans cave. South African Journal of Science 84, 828e835.
  25. Broom, R., 1938. The Pleistocene anthropoid apes of South Africa. Nature 142, 377e379.
  26. Broom, R., Robinson, J.T., 1949. Another new type of fossil ape-man. Nature 163, 57e57.
  27. Butzer, K.W., 1976. Lithostratigraphy of the Swartkrans Formation. South African Journal of Science 72, 136e141.
  28. Cadman, A., Rayner, R.J., 1989. Climatic change and the appearance of Austral- opithecus africanus in the Makapansgat sediments. Journal of Human Evolution 18, 107e113.
  29. Calandra, I., Merceron, G., 2016. Dental microwear texture analysis in mammalian ecology. Mammal Review 46, 215e228.
  30. Calandra, I., Schulz, E., Pinnow, M., Krohn, S., Kaiser, T.M., 2012. Teasing apart the contributions of hard dietary items on 3D dental microtextures in primates. Journal of Human Evolution 63, 85e98.
  31. Carri on, J.S., Scott, L., 1999. The challenge of pollen analysis in palaeoenvironmental studies of hominid beds: the record from Sterkfontein caves. Journal of Human Evolution 36, 401e408.
  32. Chivers, D.J., Wood, B.A., Bilsborough, A., 1984. Food Acquisition and Processing in Primates. Plenum, New York.
  33. Churcher, C.S., Watson, V., 1993. Additional fossil Equidae from Swartkrans. In: Brain, C.K. (Ed.), Swartkrans: A Cave's Chronicle of Early Man, Transvaal Museum Monograph No. 8. Transvaal Museum, Pretoria, pp. 137e150.
  34. Clarke, R.J., 2006. A deeper understanding of the stratigraphy of Sterkfontein fossil hominid site. Transactions of the Royal Society of South Africa 61, 111e120.
  35. Clarke, R.J., Tobias, P.V., 1995. Sterkfontein Member 2 foot bones of the oldest South African hominid. Science 269, 521e524.
  36. Codron, D., Luyt, J., Lee-Thorp, J.A., Sponheimer, M., de Ruiter, D., Codron, J., 2005. Utilization of savanna-based resources by Plio-Pleistocene baboons. South Af- rican Journal of Science 101, 245e248.
  37. Conover, W.J., Iman, R.L., 1981. Rank transformations as a bridge between para- metric and nonparametric statistics. The American Statistician 35, 124e129.
  38. Cook, R.J., Farewell, V.T., 1996. Multiplicity considerations in the design and analysis of clinical trials. Journal of the Royal Statistical Society A 159, 93e100.
  39. Daegling, D.J., McGraw, W.S., Ungar, P.S., Pampush, J.D., Vick, A.E., Bitty, E.A., 2011. Hard-object feeding in sooty mangabeys (Cercocebus atys) and interpretation of early hominin feeding ecology. PLoS One 6, e23095.
  40. Daegling, D.J., Judex, S., Ozcivici, E., Ravosa, M.J., Taylor, A.B., Grine, F.E., Teaford, M.F., Ungar, P.S., 2013. Feeding mechanics, diet and dietary adaptations in early hominins. American Journal of Physical Anthropology 151, 356e371.
  41. De Graaff, G., 1960. A preliminary investigation of the mammalian microfauna in Pleistocene deposits of caves in the Transvaal system. Palaeontologia Africana 7, 59e117.
  42. De Graaff, G., 1961. On the fossil mammalian microfauna collected at Kromdraai by Draper in 1895. South African Journal of Science 57, 259e260.
  43. Delezene, L., Teaford, M.F., Ungar, P.S., 2016. Canine and incisor microwear in pitheciids and Ateles reflects documented patterns of tooth use. American Journal of Physical Anthropology 161, 6e25.
  44. Delson, E., 1988. Chronology of South African australopith site units. In: Grine, F.E. (Ed.), Evolutionary History of the ''Robust'' Australopithecines. Aldine de Gruyter, New York, pp. 317e324. deMenocal, P.B., Tierney, J.E., 2012. Green Sahara: African humid periods paced by Earth's orbital changes. Nature Education Knowledge 3, 12.
  45. Denys, C., 1990. Implications pal eo ecologiques et pal eobiog eographiques de l' etude de rongeurs Plio-Pleistocene d'Afrique orientale et australe. Ph.D. Dissertation, Universit e Pierre et Marie Curie (Paris 6).
  46. Denys, C., 1992. Les analyses multivari ees: une aide a l'interpr etation des pal eoenvironments. L'example des rongeurs Plio-Pl eistoc ene d'Afrique australe. Geobios 14, 209e217.
  47. Denys, C., 1999. Of mice and men: evolution in East and South Africa during Plio- Pleistocene times. In: Bromage, T.G., Schrenk, F. (Eds.), African Biogeography, Climate Change and Human Evolution. Oxford University Press, Oxford, pp. 226e252.
  48. de Ruiter, D.J., 2003. Revised faunal lists for Members 1e3 of Swartkrans. Annals of the Transvaal Museum 40, 29e41.
  49. de Ruiter, D.J., 2004. Undescribed hominin fossils from the Transvaal Museum faunal collections. Annals of the Transvaal Museum 41, 29e40.
  50. de Ruiter, D.J., Sponheimer, M., Lee-Thorp, J.A., 2008. Indications of habitat associ- ation of Australopithecus robustus in the Blaaubank Valley, South Africa. Journal of Human Evolution 55, 1015e1030.
  51. DeSantis, L.R., Schubert, B.W., Scott, J.R., Ungar, P.S., 2012. Times not so tough at La Brea: dental microwear texture analysis clarifies the feeding behavior of the saber-toothed cat, Smilodon fatalis, and American lion, Panthera atrox. Journal of Vertebrate Paleontology 32, 86e87.
  52. Domínguez-Rodrigo, M., Musiba, C.M., 2010. How accurate are paleoecological re- constructions of early paleontological and archaeological sites? Evolutionary Biology 37, 128e140.
  53. Donohue, S.L., DeSantis, L.R.G., Schubert, B.W., Ungar, P.S., 2013. Was the giant short-faced bear a hyper-scavenger? A new approach to the dietary study of ursids using dental microwear textures. PLoS One 8, e77531.
  54. Elton, S., 2001. Locomotor and habitat classifications of cercopithecoid postcranial material from Sterkfontein Member 4, Bolt's Farm and Swartkrans Members 1 and 2, South Africa. Palaeontologia Africana 37, 115e126.
  55. El-Zaatari, S., 2010. Occlusal microwear texture analysis and the diets of historical/ prehistoric hunter-gatherers. International Journal of Osteoarchaeology 20, 67e87.
  56. El-Zaatari, S., Grine, F.E., Teaford, M.F., Smith, H.F., 2005. Molar microwear and di- etary reconstructions of fossil Cercopithecoidea from the Plio-Pleistocene de- posits of South Africa. Journal of Human Evolution 49, 180e205.
  57. El-Zaatari, S., Grine, F.E., Ungar, P.S., Hublin, J.J., 2011. Ecogeographic variation in Neandertal dietary habits: Evidence from occlusal molar microwear texture analysis. Journal of Human Evolution 61, 411e424.
  58. Fleagle, J.G., 2013. Primate Adaptation and Evolution, third edition. Academic Press, New York.
  59. Freedman, L., 1970. A new check-list of fossil Cercopithecoidea of South Africa. Palaeontologia Afrricana 13, 109e110.
  60. Frost, S.R., Delson, E., 2002. Fossil Cercopithecidae from the Hadar Formation and surrounding areas of the Afar Depression, Ethiopia. Journal of Human Evolution 43, 687e748.
  61. Frost, S.R., Gilbert, C.C., Pugh, K.D., Guthrie, E.H., Delson, E., 2015. The hand of Cercopithecoides williamsi (Mammalia, Primates): earliest evidence for thumb reduction among colobine monkeys. PLoS One 10, e0125030.
  62. Gallagher, A., Menter, C.G., 2011. DNH 109: a fragmentary hominin near-proximal ulna from Drimolen. South Africa. South African Journal of Science 107, 1e4.
  63. Gamza, T.R., 2010. Intra-individual microwear variation: deciduous versus perma- nent dentition. Dental Anthropology 23, 66e68.
  64. Gommery, D., Braga, J., Keyser, A., 1998. Drimolen: Des b eb es de deux millions d'ann ees. Arch eologia 343, 20e27.
  65. Gommery, D., Senut, B., Keyser, A., 2002. A fragmentary pelvis of Paranthropus robustus of the Plio-Pleistocene site of Drimolen (Republic of South Africa). Geobios 35, 265e281.
  66. Gordon, K.D., 1982. A study of microwear on chimpanzee molars: implications for dental microwear analysis. American Journal of Physical Anthropology 59, 195e215.
  67. Gordon, K.D., 1984. Hominoid dental microwear: complications in the use of microwear analysis to detect diet. Journal of Dental Research 63, 1043e1046.
  68. Grine, F.E., 1977. Analysis of early hominid deciduous molar wear by scanning electron microscopy: a preliminary report. Proceedings of the Electron Micro- scopy Society of South Africa 7, 157e158.
  69. Grine, F.E., 1981. Trophic differences between 'gracile' and 'robust' australopithe- cines: a scanning electron microscope analysis of occlusal events. South African Journal of Science 77, 203e230.
  70. Grine, F.E., 1982. Note on a new hominid specimen from Member 3, Kromdraai Formation, Transvaal. Annals of the Transvaal Museum 33, 287e290.
  71. Grine, F.E., 1984. Deciduous molar microwear of South African australopithecines. In: Chivers, D.J., Wood, B.A., Bilsborough, A. (Eds.), Food Acquisition and Pro- cessing in Primates. Plenum, New York, pp. 525e534.
  72. Grine, F.E., 1986. Dental evidence for dietary differences in Australopithecus and Paranthropus: a quantitative analysis of permanent molar microwear. Journal of Human Evolution 15, 783e822.
  73. Grine, F.E., 1989. New hominid Fossils from the Swartkrans Formation (1979e1986 excavations): craniodental specimens. American Journal of Physical Anthro- pology 79, 409e449.
  74. Grine, F.E., 2013. The alpha taxonomy of Australopithecus africanus. In: Reed, K.E., Fleagle, J.G., Leakey, R.E. (Eds.), The Paleobiology of Australopithecus. Springer, Dordrecht, pp. 73e104.
  75. Grine, F.E., Daegling, D.J., 2017. Functional morphology, biomechanics and the retrodiction of early hominin diets. Comptes Rendus Palevol 16, 613e631.
  76. Grine, F.E., Kay, R.F., 1988. Early hominid diets from quantitative image analysis of dental microwear. Nature 333, 765e768.
  77. Grine, F.E., Sponheimer, M., Ungar, P.S., Lee-Thorp, J., Teaford, M.F., 2012. Dental microwear and stable isotopes inform the paleoecology of extinct hominins. American Journal of Physical Anthropology 148, 285e317.
  78. Gügel, I.L., Grupe, G., Kunzelmann, K.H., 2001. Simulation of dental microwear: Characteristic traces by opal phytoliths give clues to ancient human dietary behavior. American Journal of Physical Anthropology 114, 124e138.
  79. Hadly, E.A., 1999. Fidelity of terrestrial vertebrate fossils to a modern ecosystem. Palaeogeography, Palaeoclimatology, Palaeoecology 149, 389e409.
  80. Haupt, R.J., DeSantis, L.R.G., Green, J.L., Ungar, P.S., 2013. Dental microwear texture as a proxy for diet in xenarthrans. Journal of Mammalogy 94, 856e866.
  81. Heaton, J.L., 2006. Taxonomy of the Sterkfontein fossil Cercopithecinae: the Papionini of Members 2 and 4 (Gauteng, South Africa). Ph.D. Dissertation, Indiana University.
  82. Hendey, Q.B., 1973. Carnivore remains from the Kromdraai australopithecine site (Mammalia: Carnivora). Annals of the Transvaal Museum 28, 99e112.
  83. Hoffman, J.M., Fraser, D., Clementz, M.T., 2015. Controlled feeding trials with un- gulates: a new application of in vivo dental molding to assess the abrasive factors of microwear. Journal of Experimental Biology 218, 1538e1547.
  84. Hopley, P.J., Maslin, M.A., 2010. Climate-averaging of terrestrial faunas: an example from the Plio-Pleistocene of South Africa. Paleobiology 36, 32e50.
  85. Hopley, P.J., Latham, A.G., Marshall, J.D., 2006. Palaeoenvironments and palaeodiets of mid-Pliocene micromammals from Makapansgat Limeworks, South Africa: A stable isotope and dental microwear approach. Palaeogeography, Palae- oclimatology, Palaeoecology 233, 235e251.
  86. Hua, L.C., Brandt, E.T., Meullenet, J.F., Zhou, Z.R., Ungar, P.S., 2015. An in vitro study of dental microwear formation using the BITE Master II chewing machine. American Journal of Physical Anthropology 158, 769e775.
  87. Kaiser, T.M., Clauss, M., Schulz-Kornas, E., 2016. A set of hypotheses on tribology of mammalian herbivore teeth. Surface Topography: Metrology and Properties 4, 014003.
  88. Karme, A., Rannikko, J., Kallonen, A., Clauss, M., Fortelius, M., 2016. Mechanical modelling of tooth wear. Journal of the Royal Society Interface 13, 20160399.
  89. Keyser, A.W., 2000. The Drimolen skull: the most complete australopithecine cra- nium and mandible to date. South African Journal of Science 96, 189e193.
  90. Keyser, A.W., Menter, C.G., Moggi-Cecchi, J., Pickering, T.R., Berger, L.R., 2000. Dri- molen: a new hominid-bearing site in Gauteng, South Africa. South African Journal of Science 96, 193e197.
  91. Kibii, J.M., 2007. Taxonomy and taphonomy of non-hominid primates from the Jacovec Cavern, Sterkfontein. South African Archaeology Bulletin 62, 90e97.
  92. Kidwell, S.M., Flessa, K.W., 1995. The quality of the fossil record: populations, species, and communities. Annual Review of Ecology and Systematics 26, 269e299.
  93. King, T., Andrews, P., Boz, B., 1999. Effect of taphonomic processes on dental microwear. American Journal of Physical Anthropology 108, 359e373.
  94. Krueger, K.L., Scott, J.R., Kay, R.F., Ungar, P.S., 2008. Dental microwear textures of "Phase I" and "Phase II" facets. American Journal of Physical Anthropology 137, 485e490.
  95. Kuman, K., Clarke, R.J., 2000. Stratigraphy, artefact industries and hominid associ- ations for Sterkfontein, Member 5. Journal of Human Evolution 38, 827e847.
  96. Leakey, M.G., 1982. Extinct large colobines from the Plio-Pleistocene of Africa. American Journal of Physical Anthropology 58, 153e172.
  97. Lee-Thorp, J.A., Sponheimer, M., 2013. Hominin ecology from hard tissue biogeo- chemistry. In: Sponheimer, M., Lee-Thorp, J.A., Reed, K.E., Ungar, P.S. (Eds.), Early Hominin Paleoecology. University of Colorado Press, Boulder, pp. 281e324.
  98. Lee-Thorp, J.A., van der Merwe, N., 1993. Stable isotope studies of Swartkrans. In: Brain, C.K. (Ed.), Swartkrans: A Cave's Chronicle of Early Man, Transvaal Museum Monograph 8. Transvaal Museum, Pretoria, pp. 251e256.
  99. Lee-Thorp, J.A., van der Merwe, N.J., Brain, C.K., 1994. Diet of Australopithecus robustus at Swartkrans from stable carbon isotopic analysis. Journal of Human Evolution 27, 361e372.
  100. Lee-Thorp, J., Thackeray, J.F., van der Merwe, N., 2000. The hunters and the hunted revisited. Journal of Human Evolution 39, 565e576.
  101. Lee-Thorp, J.A., Sponheimer, M., van der Merwe, N.J., 2003. What do stable isotopes tell us about hominid dietary and ecological niches in the Pliocene? Interna- tional Journal of Osteoarchaeology 13, 104e113.
  102. Lee-Thorp, J.A., Sponheimer, M., Luyt, J., 2007. Tracking changing environments using stable carbon isotopes in fossil tooth enamel: an example from the South African hominin sites. Journal of Human Evolution 53, 595e601.
  103. Leichliter, J.N., 2011. Micromammal paleoecology: theory, methods, and application to modern and fossil assemblages in the Cradle of Humankind World Heritage Site, South Africa. M.A. Thesis, University of Colorado.
  104. Lucas, P.W., Omar, R., Al-Fadhalah, K., Almusallam, A.S., Henry, A.G., Michael, S., Thai, L.A., Watzke, J., Strait, D.S., Atkins, A.G., 2013. Mechanisms and causes of wear in tooth enamel: implications for hominin diets. Journal of the Royal Society Interface 10, 20120923.
  105. Lucas, P.W., van Casteren, A., Al-Fadhalah, K., Almusallam, A.S., Henry, A.G., Michael, S., Watzke, J., Reed, D.A., Diekwisch, T.G.H., Strait, D.S., Atkins, A.G., 2014. The role of dust, grit and phytoliths in tooth wear. Annales Zoologici Fennici 51, 143e152.
  106. Luyt, J., Lee-Thorp, J.A., 2003. Carbon isotope ratios of Sterkfontein fossils indicate a marked shift to open environments ca. 1.7 Ma. South African Journal of Science 99, 271e273.
  107. Lyman, R.L., 2017. Paleoenvironmental reconstruction from faunal remains: ecological basics and analytical assumptions. Journal of Archaeological Research 25, 315e371.
  108. Mahoney, P., 2006a. Intertooth and intrafacet dental microwear variation in an archaeological sample of modern humans from the Jordan Valley. American Journal of Physical Anthropology 129, 39e44.
  109. Mahoney, P., 2006b. Dental microwear from Natufian hunter-gatherers and early Neolithic farmers: comparisons within and between samples. American Journal of Physical Anthropology 130, 308e319.
  110. Mainland, I.L., 1998. Dental microwear and diet in domestic sheep (Ovis aries) and goats (Capra hircus): distinguishing grazing and fodder-fed ovicaprids using a quantitative analytical approach. Journal of Archaeological Science 25, 1259e1271.
  111. Mainland, I.L., 2003. Dental microwear in grazing and browsing Gotland sheep (Ovis aries) and its implications for dietary reconstruction. Journal of Archaeological Science 30, 1513e1527.
  112. McKee, J.K., 1991. Palaeo-ecology of the Sterkfontein hominids: A review and syn- thesis. Palaeontologia Africana 28, 41e51.
  113. McKee, J.K., 1999. The autocatalytic nature of hominid evolution in African Plio- Pleistocene environments. In: Bromage, T.G., Schrenk, F. (Eds.), African Bioge- ography, Climate Change, and Human Evolution. Oxford University Press, Ox- ford, pp. 57e75.
  114. Mendelsohn, J.P., 1989. Habitat preferences, population size, food and breeding of six owl species in the Springbok Flats, South Africa. Ostrich 60, 183e190.
  115. Merceron, G., Escarguel, G., Angibault, J.M., Verheyden-Tixier, H., 2010. Can dental microwear textures record inter-individual dietary variations? PLoS One 5 e9542. Merceron, G., Ramdarshan, A., Blondel, C., Boisserie, J.-R., Brunetiere, N., Francisco, A., Gautier, D., Milhet, X., Novello, A., Pret, D., 2016. Untangling the environmental from the dietary: dust does not matter. Proceedings of the Royal Society B 283, 20161032.
  116. Merceron, G., Ramdarshan, A., Blondel, C., Boisserie, J.-R., Brunetiere, N., Francisco, A., Gautier, D., Milhet, X., Novello, A., Pret, D., Surault, J., 2017. Food, dust and tooth wear: a sheep perspective. In: 17th International Symposium on Dental Morphology & 2nd Congress of International Association for Paleo- dontology, p. 109.
  117. Moggi-Cecchi, J., Grine, F.E., Tobias, P.V., 2006. Early hominid dental remains from Members 4 and 5 of the Sterkfontein Formation (1966e1996 excavations): catalogue, individual associations, morphological descriptions and initial metrical analysis. Journal of Human Evolution 50, 239e328.
  118. Moggi-Cecchi, J., Menter, C., Boccone, S., Keyser, A., 2010. Early hominin dental remains from the Plio-Pleistocene site of Drimolen, South Africa. Journal of Human Evolution 58, 374e405.
  119. O'Regan, H.J., Menter, C.G., 2009. Carnivora from the Plio-Pleistocene hominin site of Drimolen, Gauteng, South Africa. Geobios 42, 329e350.
  120. O'Regan, H.J., Reynolds, S.C., 2009. An ecological reassessment of the southern Af- rican carnivore guild: a case study from Member 4, Sterkfontein, South Africa. Journal of Human Evolution 57, 212e222.
  121. Partridge, T.C., 1978. Re-appraisal of the lithostratigraphy of Sterkfontein hominid site. Nature 275, 282e287.
  122. Partridge, T.C., 1979. Re-appraisal of the lithostratigraphy of Makapansgat Lime- works hominid site. Nature 279, 484e488.
  123. Partridge, T.C., 1982. Some preliminary observations on the stratigraphy and sedi- mentology of the Kromdraai B hominid site. In: Coetzee, J.A., Van Zinderen Bakker, E.M. (Eds.), Palaeoecology of Africa and the Surrounding Islands, vol. 15. CRC Press, New York, pp. 3e12.
  124. Partridge, T.C., 1985. The palaeoclimatic significance of Cainozoic terrestrial strati- graphic and tectonic evidence from Southern Africa: a review. South African Journal of Science 81, 245e247.
  125. Partridge, T.C., 2000. Hominid-bearing cave and tufa deposits. In: Partridge, T.C., Maud, R.R. (Eds.), The Cenozoic in Southern Africa. Oxford University Press, Oxford, pp. 100e130.
  126. Partridge, T.C., Watt, I.B., 1991. The stratigraphy of the Sterkfontein hominid deposit and its relationship to the underground cave system. Palaeontologia Africana 28, 35e40.
  127. Partridge, T.C., Granger, D.E., Caffee, M.W., Clarke, R.J., 2003. Lower Pliocene hom- inid remains from Sterkfontein. Science 300, 607e612.
  128. Percher, A.M., Merceron, G., Akoue, G.N., Galbany, J., Romero, A., Charpentier, M.J.E., 2017. Dental microwear textural analysis as an analytical tool to depict indi- vidual traits and reconstruct the diet of a primate. American Journal of Physical Anthropology 165, 123e138.
  129. Perrin, M.R., 2011. A fossilised humerus of a lovebird tells little of the Pleistocene habitat of Australopithecus robustus. South African Journal of Science 107, 9e10. Petersen, C.H., 1977. The paleoecological significance of undetected short-term variability. Journal of Paleontology 51, 976e981.
  130. Plavcan, J.M., Cope, D.A., 2001. Metric variation and species recognition in the fossil record. Evolutionary Anthropology 10, 204e222.
  131. Pocock, T.N., 1971. Pleistocene bird fossils from Kromdraai and Sterkfontein. Ostrich 8, 1e6.
  132. Pocock, T.N., 1987. Plio-Pleistocene mammalian microfauna in southern Africa, a preliminary report including description of two new fossil muroid genera (Mammalia: Rodentia). Palaeontologia Africana 26, 69e91.
  133. Pontzer, H., Scott, J.R., Lordkipanidze, D., Ungar, P.S., 2011. Dental microwear texture analysis and diet in the Dmanisi hominins. Journal of Human Evolution 61, 683e687.
  134. Prideaux, G.J., Ayliffe, L.K., DeSantis, L.R.G., Schubert, B.W., Murray, P.F., Gagan, M.K., Cerling, T.E., 2009. Extinction implications of a chenopod browse diet for a giant Pleistocene kangaroo. Proceedings of the National Academy of Sciences USA 106, 11646e11650.
  135. Puech, P.F., Prone, A., 1979. Reproduction exp erimentale des processes d'usure dentaire par abrasion: implications pal eoecologique chez l'Homme fossile. Comptes Rendus Hebdomadaires des Seances de l'Academie des Sciences Serie D 289, 895e898.
  136. Purnell, M.A., Crumpton, N., Gill, P.G., Jones, G., Rayfield, E.J., 2013. Within-guild dietary discrimination from 3-D textural analysis of tooth microwear in insectivorous mammals. Journal of Zoology 291, 249e257.
  137. Ragni, A.J., Teaford, M.F., Ungar, P.S., 2017. A molar microwear texture analysis of pitheciid primates. American Journal of Primatology 79, e22697.
  138. Ramdarshan, A., Blondel, C., Brunetiere, N., Francisco, A., Gautier, D., Surault, J., Merceron, G., 2016. Seeds, browse, and tooth wear: a sheep perspective. Ecol- ogy and Evolution 6, 5559e5569.
  139. Ramdarshan, A., Blondel, C., Gautier, D., Brunetiere, N., Surault, J., Merceron, G., 2017. Overcoming sampling issues in dental tribology: Insights from an experimentation on sheep. Palaeontologia Electronica 20, 53A.
  140. Rayner, R.J., Moon, B.P., Masters, J.C., 1993. The Makapansgat australopithecine environment. Journal of Human Evolution 24, 219e231.
  141. Reed, D.N., 2005. Taphonomic implications of roosting behavior and trophic habits in two species of African owl. Journal of Archaeological Science 32, 1669e1676.
  142. Reed, K., 1997. Early hominid evolution and ecological change through the African Plio-Pleistocene. Journal of Human Evolution 32, 289e322.
  143. Reed, K., 1998. Using large mammal communities to examine ecological and taxonomic organization and predict vegetation in extant and extinct assem- blages. Paleobiology 32, 384e408.
  144. Reed, K.E., Rector, A.L., 2006. African Pliocene paleoecology: hominin habitats, re- sources, and diets. In: Ungar, P.S. (Ed.), Evolution of the Human Diet: The Known, the Unknown, and the Unknowable. Oxford University Press, Oxford, pp. 262e288.
  145. Reynolds, S.C., Kibii, J.M., 2011. Sterkfontein at 75: review of palaeoenvironments, fauna, and archaeology from the hominin site of Sterkfontein (Gauteng Prov- ince, South Africa). Palaeontologia Africana 46, 59e88.
  146. Reynolds, S.C., Vogel, J.C., Clarke, R.J., Kuman, K.A., 2003. Preliminary results of excavations at Lincoln Cave, Sterkfontein. South African Journal of Science 99, 286e288.
  147. Reynolds, S.C., Clarke, R.J., Kuman, K.A., 2007. The view from the Lincoln Cave: Mid- to Late Pleistocene fossil deposits from the Sterkfontein hominid site, South Africa. Journal of Human Evolution 53, 260e271.
  148. Reynolds, S.C., Bailey, G.N., King, G.C.P., 2011. Landscapes and their relation to hominin habitats: case studies from Australopithecus sites in eastern and southern Africa. Journal of Human Evolution 60, 281e298.
  149. Rivals, F., Semprebon, G.M., 2006. A comparison of the dietary habits of a large sample of the Pleistocene pronghorn Stockoceros onusrosagris from the Papago Springs Cave in Arizona to the modern Antilocapra americana. Journal of Vertebrate Paleontology 26, 495e500.
  150. Robinson, J.T., 1952. The australopithecine-bearing deposits of the Sterkfontein area. Annals of the Transvaal Museum 22, 1e19.
  151. Robinson, J.T., 1962. Australopithecines and artefacts at Sterkfontein. Part I. Sterk- fontein stratigraphy and the significance of the Extension Site. South African Archaeological Bulletin 17, 87e107.
  152. Robson, S.K., Young, W.G., 1986. Tooth microwear of Thylacinus cynocephalus and Sarcophilus harrisii. Journal of Dental Research 65, 483e483.
  153. Ross, C.F., Iriarte-Diaz, J., 2014. What does feeding system morphology tell us about feeding? Evolutionary Anthropology 23, 105e120.
  154. Rovinsky, D.S., Herries, A.I.R., Menter, C.G., Adams, J.W., 2015. First description of in situ primate and faunal remains from the Plio-Pleistocene Drimolen Makondo palaeocave infill, Gauteng, South Africa. Palaeontologia Electronica 18, 34A.
  155. Roy, K., Valentine, J.W., Jablonski, D., Kidwell, S.M., 1996. Scales of climatic vari- ability and time averaging in Pleistocene biotas: implications for ecology and evolution. Trends in Ecology & Evolution 11, 458e463.
  156. Sanson, G.D., Kerr, S.A., Gross, K.A., 2007. Do silica phytoliths really wear mammalian teeth? Journal of Archaeological Science 34, 526e531.
  157. Schubert, B.W., 2007. Dental mesowear and the palaeodiets of bovids from Maka- pansgat Limeworks Cave, South Africa. Palaeontologia Africana 42, 43e50.
  158. Schubert, B.W., Ungar, P.S., Sponheimer, M., Reed, K.E., 2006. Microwear evidence for Plio-Pleistocene bovid diets from Makapansgat Limeworks Cave, South Af- rica. Palaeogeography, Palaeoclimatology, Palaeoecology 241, 301e319.
  159. Schubert, B.W., Ungar, P.S., DeSantis, L.R.G., 2010. Carnassial microwear and dietary behaviour in large carnivorans. Journal of Zoology 280, 257e263.
  160. Schulz, E., Piotrowski, V., Clauss, M., Mau, M., Merceron, G., Kaiser, T.M., 2013. Di- etary abrasiveness is associated with variability of microwear and dental sur- face texture in rabbits. PLoS One 8, e56167.
  161. Scott, J.R., 2012. Dental microwear texture analysis of extant African Bovidae. Mammalia 76, 157e174.
  162. Scott, J.R., Godfrey, L.R., Jungers, W.L., Scott, R.S., Simons, E.L., Teaford, M.F., Ungar, P.S., Walker, A., 2009. Dental microwear texture analysis of two families of subfossil lemurs from Madagascar. Journal of Human Evolution 56, 405e416.
  163. Scott, L., 1982. Pollen analyses of Late Cainozoic deposits in the Transvaal, South Africa, and their bearing on palaeoclimates. Palaeoecology 15, 101e107.
  164. Scott, L., 1995. Pollen evidence for vegetational and climatic change in southern Africa during the Neogene and Quaternary. In: Vrba, E.S., Denton, G.H., Partridge, T.C., Burckle, L.H. (Eds.), Paleoclimate and Evolution with Emphasis on Human Origins. Yale University Press, New Haven, pp. 65e76.
  165. Scott, L., Bonnefille, R., 1986. Search for pollen from the hominid deposits of Kromdraai, Sterkfontein and Swartkrans: some problems and preliminary re- sults. South African Journal of Science 82, 380e382.
  166. Scott, R.S., Ungar, P.S., Bergstrom, T.S., Brown, C.A., Grine, F.E., Teaford, M.F., Walker, A., 2005. Dental microwear texture analysis shows within-species diet variability in fossil hominins. Nature 436, 693e695.
  167. Scott, R.S., Ungar, P.S., Bergstrom, T.S., Brown, C.A., Childs, B.E., Teaford, M.F., Walker, A., 2006. Dental microwear texture analysis: Technical considerations. Journal of Human Evolution 51, 339e349.
  168. Scott, R.S., Teaford, M.F., Ungar, P.S., 2012. Dental microwear texture and anthropoid diets. American Journal of Physical Anthropology 147, 551e579.
  169. S en egas, F., Paradis, E., Michaux, J., 2005. Homogeneity of fossil assemblages extracted from mine dumps: an analysis of Plio-Pleistocene fauna from South African caves. Lethaia 38, 315e322.
  170. Silcox, M.T., Teaford, M.F., 2002. The diet of worms: An analysis of mole dental microwear. Journal of Mammalogy 83, 804e814.
  171. Skinner, J.D., Chimimba, C.T., 2005. The Mammals of the Southern African Subre- gion, third edition. Cambridge University Press, Cambridge.
  172. Sponheimer, M., 1999. Isotopic ecology of the Makapansgat Limeworks fauna. Ph.D. Dissertation, Rutgers University.
  173. Sponheimer, M., Lee-Thorp, J.A., 1999. Isotopic evidence for the diet of an early hominid, Australopithecus africanus. Science 283, 368e370.
  174. Sponheimer, M., Lee-Thorp, J.A., 2003. Using bovid carbon isotope data to provide palaeoenvironmental information. South African Journal of Science 99, 273e275.
  175. Sponheimer, M., Lee-Thorp, J.A., 2009. Biogeochemical evidence for the environ- ments of early Homo in South Africa. In: Grine, F.E., Fleagle, J.G., Leakey, R.E. (Eds.), The First Humans. Origin and Early Evolution of the Genus Homo. Springer, Dordrecht, pp. 185e194.
  176. Sponheimer, M., Reed, K.E., Lee-Thorp, J.A., 1999. Combining isotopic and ecomor- phological data to refine bovid palaeodietary reconstruction: a case study from Makapansgat Limeworks hominin locality. Journal of Human Evolution 36, 705e718.
  177. Sponheimer, M., Reed, K.E., Lee-Thorp, J.A., 2001. Isotopic palaeoecology of Maka- pansgat Limeworks Perissodactyla. South African Journal of Science 97, 327e329.
  178. Steininger, C.M., 2011. The dietary behaviour of Early Pleistocene bovids from Cooper's Cave and Swartkrans, South Africa. Ph.D. Dissertation, University of the Witwatersrand.
  179. Stidham, T.A., 2009. A lovebird (Psittaciformes: Agapornis) from the Plio-Pleistocene Kromdraai B locality, South Africa. South African Journal of Science 105, 155e157.
  180. Strait, S.G., 1993. Molar morphology and food texture among small bodied insec- tivorous mammals. Journal of Mammalogy 74, 391e402.
  181. Strait, D.S., Constantino, P., Lucas, P.W., Richmond, B.G., Spencer, M.A., Dechow, P.C., Ross, C.F., Grosse, I.R., Wright, B.W., Wood, B.A., Weber, G.W., Wang, Q., Byron, C., Slice, D.E., Chalk, J., Smith, A.L., Smith, L.C., Wood, S., Berthaume, M., Benazzi, S., Dzialo, C., Tamvada, K., Ledogar, J.A., 2013. Diet and dietary adap- tations in early hominins: the hard food perspective. American Journal of Physical Anthropology 151, 339e355.
  182. Stratford, D., Heaton, J.L., Pickering, T.R., Caruana, M.V., Shadrach, K., 2016. First hominin fossils from Milner Hall, Sterkfontein, South Africa. Journal of Human Evolution 91, 167e173.
  183. Strier, K.B., 2016. Primate Behavior Ecology, 5th Edition. Routledge, New York.
  184. Stynder, D.D., Ungar, P.S., Scott, J.R., Schubert, B.W., 2012. A dental microwear texture analysis of the Mio-Pliocene hyaenids from Langebaanweg, South Af- rica. Acta Palaeontologica Polonica 57, 485e496.
  185. T ¸ alu, S., Stach, S., Mahajan, A., Pathak, D., Wagner, T., Kumar, A., Bedi, R.K., 2014. Multifractal analysis of drop-casted copper (II) tetrasulfophthalocyanine film surfaces on the indium tin oxide substrates. Surface and Interface Analysis 46, 393e398.
  186. Teaford, M.F., 1985. Molar microwear and diet in the genus Cebus. American Journal of Physical Anthropology 66, 363e370.
  187. Teaford, M.F., 1988. Scanning electron microscope diagnosis of wear patterns versus artifacts on fossil teeth. Scanning Microscopy 2, 1167e1175.
  188. Teaford, M.F., Lytle, J.D., 1996. Diet-induced changes in rates of human tooth microwear: A case study involving stone-ground maize. American Journal of Physical Anthropology 100, 143e147.
  189. Teaford, M.F., Oyen, O.J., 1989. Dental microwear in vervets raised on different diets. Journal of Dental Research 65, 278e278.
  190. Teaford, M.F., Walker, A., 1983. Dental microwear in adult and still-born guinea pigs (Cavia porcellus). Archives of Oral Biology 28, 1077e1081.
  191. Teaford, M.F., Walker, A., 1984. Quantitative differences in dental microwear be- tween primate species with different diets and a comment on the presumed diet of Sivapithecus. American Journal of Physical Anthropology 64, 191e200.
  192. Turner, A., 1986. Miscellaneous carnivore remains from Plio-Pleistocene deposits in the Sterkfontein Valley (Mammalia: Carnivora). Annals of the Transvaal Museum 34, 205e225.
  193. Ungar, P.S., Hlusko, L.J., 2016. The evolutionary path of least resistance. Science 353, 29e30.
  194. Ungar, P.S., Scott, R.S., 2009. Dental evidence for diets of Early Homo. In: Grine, F.E., Leakey, R.E., Fleagle, J.G. (Eds.), The First Humans: Origins of the Genus Homo. Springer-Verlag, New York, pp. 121e134.
  195. Ungar, P.S., Merceron, G., Scott, R.S., 2007. Dental microwear texture analysis of Vars- water bovids and early Pliocene paleoenvironments of Langebaanweg, Western Cape Province, South Africa. Journal of Mammalian Evolution 14, 163e181.
  196. Ungar, P.S., Grine, F.E., Teaford, M.F., 2008. Dental microwear and diet of the Plio- Pleistocene hominin Paranthropus boisei. PLoS One 3 e2044.
  197. Ungar, P.S., Scott, J.R., Schubert, B.W., Stynder, D.D., 2010a. Carnivoran dental microwear textures: comparability of carnassial facets and functional differ- entiation of postcanine teeth. Mammalia 74, 219e224.
  198. Ungar, P.S., Scott, R.S., Grine, F.E., Teaford, M.F., 2010b. Molar microwear textures and the diets of Australopithecus anamensis and Australopithecus afarensis. Philosophical Transactions of the Royal Society 365, 3345e3354.
  199. Ungar, P.S., Krueger, K.L., Blumenschine, R.J., Njau, J., Scott, R.S., 2012. Dental micro- wear texture analysis of hominins recovered by the Olduvai Landscape Paleo- anthropology Project, 1995e2007. Journal of Human Evolution 63, 429e437.
  200. Ungar, P.S., Scott, J.R., Steininger, C.M., 2016. Dental microwear differences between eastern and southern African fossil bovids and hominins. South African Journal of Science 112. #2015-0393.
  201. van der Merwe, N.J., Thackeray, J.F., Lee-Thorp, J.A., Luyt, J., 2003. The carbon isotope ecology and diet of Australopithecus africanus at Sterkfontein, South Africa. Journal of Human Evolution 44, 581e597.
  202. Van Valkenburgh, B., Teaford, M.F., Walker, A., 1990. Molar microwear and diet in large carnivores: Inferences concerning diet in the sabretooth cat, Smilodon fatalis. Journal of Zoology 222, 319e340.
  203. Vernon, D.S., 2013. A morphometric analysis of the phalanges and a fragmentary first metatarsal from the Drimolen hominin site, South Africa. Ph.D. Disserta- tion, University of Johannesburg.
  204. Vrba, E.S., 1974. Chronological and ecological implications of the fossil Bovidae at the Sterkfontein australopithecine site. Nature 250, 19e23.
  205. Vrba, E.S., 1975. Some evidence of chronology and palaeoecology of Sterkfontein, Swartkrans and Kromdraai from the fossil Bovidae. Nature 254, 301e304.
  206. Vrba, E.S., 1976. The Fossil Bovidae of Sterkfontein, Swartkrans and Kromdraai. Transvaal Museum Memoir No. 21. Transvaal Museum, Pretoria.
  207. Vrba, E., 1980. The significance of bovid remains as indicators of environment and predation patterns. In: Behrensmeyer, A.K., Hill, A.P. (Eds.), Fossils in the Mak- ing. University of Chicago Press, Chicago, pp. 247e272.
  208. Vrba, E.S., 1981. The Kromdraai australopithecine site revisited in 1980: recent in- vestigations and results. Annals of the Transvaal Museum 33, 17e60.
  209. Vrba, E.S., 1985a. Early hominids in southern Africa: updated observations on chronological and ecological background. In: Tobias, P.V. (Ed.), Hominid Evo- lution: Past, Present, and Future. Alan, R. Liss, New York, pp. 195e200.
  210. Vrba, E.S., 1985b. Ecological and adaptive changes associated with early hominid evolution. In: Delson, E. (Ed.), Ancestors: The Hard Evidence. Alan R. Liss, New York, pp. 63e71.
  211. Vrba, E.S., Panagos, D.C., 1982. New perspectives on taphonomy, palaeoecology and chronology of the Kromdraai ape-man. In: Coetzee, J.A., Van Zinderen Bakker, E.M. (Eds.), Palaeoecology of Africa and the Surrounding Islands, vol. 15. CRC Press, New York, pp. 13e26.
  212. Ward, J., Mainland, I.L., 1999. Microwear in modern rooting and stall-fed pigs: The potential of dental microwear analysis for exploring pig diet and management in the past. Environmental Archaeology 4, 25e32.
  213. Watson, V., 1993. Composition of the Swartkrans bone accumulations, in terms of skeletal parts and animals represented. In: Brain, C.K. (Ed.), Swartkrans: A Cave's Chronicle of Early Man, Transvaal Museum Monograph No. 8. Transvaal Museum, Pretoria, pp. 35e73.
  214. Wells, L.H., Cooke, H.B.S., 1956. Fossil Bovidae from the Limeworks quarry, Maka- pansgat, Potgietersrus. Palaeontologia Africana 4, 1e55.
  215. Wilkinson, M.J., 1985. Lower-lying and possibly older fossiliferous deposits at Sterkfontein. In: Tobias, P.V. (Ed.), Hominid Evolution: Past, Present and Future. Alan R. Liss, New York, pp. 165e170.
  216. Zavada, M.S., Cadman, A., 1993. Palynological investigations at the Makapansgat Limeworks: an australopithecine site. Journal of Human Evolution 25, 337e350.