23. I discuss this method of walking on trees by orang-utans in Humans Who Went Extinct, 35.
CHAPTER 3
1. Barbary macaques Macaca sylvanus, the famous rock apes, roam freely on Gibraltar. They were introduced from North Africa in the 18th century by the British. They live in troops and defend territories. The Gibraltar Government provides food and water for them in fixed locations but groups splinter and separate away from these sources, often raiding houses in the town below. They also forage on wild foods. During the summer in particular, these rogue groups do not have access to fixed water points.
2. Australopithecines, the species of early hominids classified under the genera Australopithecus and Paranthropus, that lived in Africa between 4.2 and 1.4 myr. I refer to them as proto-humans in The Humans Who Went Extinct.
3. The Humans Who Went Extinct, table 1, p. 26.
4. To the species listed in table 1 in The Humans Who Went Extinct we must add the new species—Australopithecus sediba—from Malapa in South Africa; R. Pickering et al., ‘Australopithecus sediba at 1.977 Ma and Implications for the Origins of the Genus Homo’, Science 333 (2011): 1421–3. In this book I will refer to this species as sediba.
5. The summaries are K. Reed, ‘Early Hominid Evolution and Ecological Change through the African Plio-Pleistocene’, Journal of Human Evolution 32 (1997): 289–322, and B. Wood and D. Strait, ‘Patterns of Resource Use in Early Homo and Paranthropus’, Journal of Human Evolution 46 (2004): 119–62.
6. Grasslands that are partially flooded or subject to regular flooding.
7. A good recent summary is P. Ungar and M. Sponheimer, ‘The Diets of Early Hominins’, Science 334 (2011): 190–3. Dental microwear patterns—usually pits and scratches in varying combinations—are detected in the teeth of fossil hominids. They reflect what an individual was eating weeks before its death, the ‘Last Supper’ phenomenon. Hard and brittle foods, such as nuts, produce dental microwear patterns that are dominated by pits and complex patterns on the tooth surface. Soft but tough foods, such as leaves, produce scratched surfaces which are typically seen as parallel lines. Surface patterns are relatively simple. Combinations of the two patterns among individuals of the same species would reflect catholic food choice relative to specialized species. Stable isotopes, especially carbon isotopes, are represented in the growth layers of teeth and reflect diet through time. In studies of early hominids they have been especially useful in identifying the proportion of C3 and C4 plants consumed. See also Ch. 2 n. 12.
8. Summarized in Ungar and Sonheimer, ‘The Diets of Early Hominins’, with additonal summary and data on two individuals of A. sediba in A. G. Henry et al., ‘The Diet of Australopithecus sediba’, Nature 487 (2012): 90–3.
9. F. E. Grine et al., ‘Molar Microwear in Praeanthropus afarensis: Evidence for Dietary Stasis through Time and under Diverse Paleoecological Conditions’, Journal of Human Evolution 51 (2006): 297–319.
10. Ungar and Sonheimer, ‘The Diets of Early Hominins’; Henry et al., ‘The Diet of Australopithecus sediba’; Grine et al., ‘Molar Microwear in Praeanthropus afarensis’.
11. Henry et al., ‘The Diet of Australopithecus sediba’.
12. From the Greek, meaning ‘plant stone’. Plants take up silica from the soil which is then deposited in various parts of the plant—the phytoliths. They are tough microscopic structures which can be preserved. The study of plant phytoliths is not new but finding them in the teeth of Neanderthals was a first for science (A. G. Henry et al., ‘Microfossils in Calculus Demonstrate Consumption of Plants and Cooked Foods in Neanderthal diets (Shanidar III, Iraq; Spy I and II, Belgium)’, Proceedings of the National Academy of Sciences USA 108 (2011): 486–91). Amanda Henry’s work then looked at earlier hominids and she recovered 38 phytoliths from Sediba.
13. M. Sponheimer et al., ‘Hominins, Sedges, and Termites: New Carbon Isotope Data from the Sterkfontein Valley and Kruger National Park’, Journal of Human Evolution 48 (2005): 301–12.
14. White et al., ‘Ardipithecus ramidus and the Paleobiology of Early Hominids’; B. Latimer and C. O. Lovejoy, ‘Hallucal Tarsometa-tarsal Joint in Australopithecus afarensis’, American Journal of Physical Anthropology 82 (1990): 125–33.
15. K. Wong, ‘Footprints to Fill: Flat Feet and Doubts about Makers of the Laetoli Tracks’, Scientific American 293 (2005): 18–19.
16. T. L. Kivell et al., ‘Australopithecus sediba Hand Demonstrates Mosaic Evolution of Locomotor and Manipulative Abilities’, Science 333 (2011): 1411–17; B. Zipfel et al., ‘The Foot and Ankle of Australopithecus sediba’, Science 333 (2011): 1417–20.
17. The Humans Who Went Extinct, table 1, p. 26.
18. Seventeen of the 21 sites (80.9 per cent) had trees and 19 of the 21 sites (90.5 per cent) had open spaces. In contrast, only 10 (47.6 per cent) had bushes.
19. Reed, ‘Early Hominid Evolution and Ecological Change’.
20. Ungar and Sponheimer, ‘The Diets of Early Hominins’. See also Ch. 2 n. 12.
21. B. Wood and M. Collard, ‘The Human Genus’, Science 284 (1999): 65–71.
22. Eight of nine sites record water, eight of nine trees, and all nine open spaces.
23. P. deMenocal, ‘African Climate Change and Faunal Evolution during the Pliocene-Pleistocene’, Earth and Planetary Science Letters 220 (2004): 3–24.
24. T. E. Cerling et al., ‘Woody Cover and Hominin Environments in the Past 6 million Years’, Nature 476 (2011): 51–6.
25. T. E. Cerling, ‘Development of Grasslands and Savannas in East Africa during the Neogene’, Palaeogeography, Palaeoclimatology, Palaeoecology 97 (1992): 241–7.
26. S. C. Reynolds, G. N. Bailey, and G. C. P. King, ‘Landscapes and Their Relation to Hominin Habitats: Case Studies from Australopithecus Sites in Eastern and Southern Africa’, Journal of Human Evolution 60 (2011): 281–98.
27. S. Semaw et al., ‘2.6 Million-Year-Old Stone Tools and Associated Bones from OGS-6 and OGS-7, Gona, Afar, Ethiopia’, Journal of Human Evolution 45 (2003): 169–77. See also Humans Who Went Extinct, 41 and 226 n. 28.
28. M. Haslam et al., ‘Primate Archaeology’, Nature 460 (2009): 339–44.
29. S. P. McPherron et al., ‘Evidence for Stone-Tool-Assisted Consumption of Animal Tissues before 3.39 Million Years Ago at Dikika, Ethiopia’, Nature 466 (2010): 857–60.
30. P. S. Ungar (ed.), Evolution of the Human Diet: The Known, the Unknown and the Unknowable (Oxford: Oxford University Press, 2007).
CHAPTER 4
1. F. Brown et al., ‘Early Homo erectus Skeleton from West Lake Turkana, Kenya’, Nature 316 (1985): 788–92.
2. Finlayson, The Humans Who Went Extinct, 54.
3. Homo erectus always incorporated trees, open spaces, and water (all seven sites examined). Bushland was only present in three (42.9 per cent) sites and rocky habitats in two (28.6 per cent). The proportions are remarkably similar to those for the australopithecines.
4. deMenocal, ‘African Climate Change’.
5. R. R. Graves et al., ‘Just How Strapping was KNM-WT 15,000?’, Journal of Human Evolution 59 (2010): 542–54.
6. H. M. McHenry, ‘Femoral Length and Stature in Plio-Pleistocene Hominids’, American Journal of Physical Anthropology 85 (1991): 149–58.
7. Estimated body weight of Homo erectus: 63 kg (139 lb, male) and 52 kg (114 lb, female). Compare with habilis: 52 kg (114 lb, male) and 32 kg (70 lb, female). From H. M. McHenry, ‘Behavioral Ecological Implications of Early Hominid Body Size’, Journal of Human Evolution 27 (1994): 77–87.
8. Energy expended per unit weight per distance moved.
9. R. H. Peters, The Ecological Implications of Body Size (Cambridge: Cambridge University Press, 1983).
10. K. L. Steudel-Numbers, ‘Energetics in Homo erectus and Other Early Hominins: The Consequences of Increased Lower-Limb Length’, Journal of Human Evolution 51 (2006): 445–53.
11. K. L. Steudel-Numbers and M. J. Tilkens, ‘The Effect of Lower Limb Length on the Energetic Cost of Locomot
ion: Implications for Fossil Hominins’, Journal of Human Evolution 47 (2004): 95–109.
12. K. L. Steudel-Numbers et al., ‘The Evolution of Human Running: Effects of Changes in Lower-Limb Length on Locomotor Economy’, Journal of Human Evolution 53 (2007): 191–6.
13. Antelopes, horses, elephants, rhinos. See, for example, T. G. Bromage and F. Schrenk (eds.), African Biogeography, Climate Change, & Human Evolution (New York: Oxford University Press, 1999).
14. Cats, dogs, and hyenas. See A. Turner, ‘The Evolution of the Guild of Larger Terrestrial Carnivores during the Plio-Pleistocene in Africa’, Geobios 23 (1990): 349–68; L. Werdelin and M. E. Lewis, ‘Plio-Pleistocene Carnivora of Eastern Africa: Species Richness and Turnover Patterns’, Zoological Journal of the Linnean Society 144 (2005): 121–44.
15. B. R. Benefit, ‘Biogeography, Dietary Specialization, and the Diversification of African Plio-Pleistocene Monkeys’, in Bromage and Schrenk (eds.), African Biogeography, 172–88.
16. T. D. White, ‘African Omnivores: Global Climatic Change and Plio-Pleistocene Hominids and Suids’, in E. S. Vrba et al. (eds.), Paleoclimate and Evolution with Emphasis on Human Origins (New Haven: Yale University Press, 1995), 369–84.
17. In southern Europe at least, wild boar Sus scrofa scavenge carcasses in a similar fashion to hyenas. E. Bernáldez Sánchez, ‘Biostratinomy Applied to the Interpretation of Scavenger Activity in Paleoecosystems’, Quaternary International 243 (2011): 161–70.
18. J. M. Baldwin, ‘A New Factor in Evolution’, American Naturalist 30 (1896): 441–51 and 536–53; but the ideas were independently conceived by Lloyd-Morgan and Osborn: C. Lloyd Morgan, ‘On Modification and Variation’, Science 4 (1896): 733–40; H. F. Osborn, ‘Ontogenic and Phylogenic Variation’, Science 4 (1896): 786–9.
19. P. Bateson, ‘The Active Role of Behaviour in Evolution’, Biology and Philosophy 19 (2004): 283–98.
20. For examples of predator activity patterns see M. Odden and P. Wegge, ‘Spacing and Activity Patterns of Leopards Panthera pardus in the Royal Bardia National Park, Nepal’, Wildlife Biology 11 (2005): 145–52; M. W. Hayward and G. Hayward, ‘Activity Patterns of Reintroduced Lion Panthera leo and Spotted Hyaena Crocuta crocuta in the Addo Elephant National Park, South Africa’, African Journal of Ecology 45 (2007): 135–41.
21. K. Modal et al., ‘Response of Leopards to Re-introduced Tigers in Sariska Tiger Reserve, Western India’, International Journal of Biodiversity and Conservation 45 (2012): 228–36.
22. Specifically cheetahs and wild dogs, B. C. R. Bertram, ‘Serengeti Predators and Their Social Systems’, in A. R. E. Sinclair and M. Norton-Griffiths (eds.), Serengeti: Dynamics of an Ecosystem (Chicago: University of Chicago Press, 1979).
23. S. M. Durant, ‘Competition Refuges and Coexistence: An Example from Serengeti Carnivores’, Journal of Animal Ecology 67 (1998): 370–86.
24. R. W. Newman, ‘Why Man Is Such a Sweaty and Thirsty Naked Animal: A Speculative Review’, Human Biology 42 (1970): 12–27.
25. For some time, the first appearance of the Acheulian and its characteristic hand-axes had been dated to 300 thousand years (hereafter kyr) after the appearance of Homo erectus. Two new papers have revised these dates and have shown that the emergence of Homo erectus and the appearance of the Acheulian were concurrent around 1.7 myr. C. J. Lepre et al., ‘An Earlier Origin for the Acheulian’, Nature 477 (2011): 82–5; Y. Beyene, ‘The Characteristics and Chronology of the Earliest Acheulean at Konso, Ethiopia’, Proceedings of the National Academy of Sciences USA 110 (2013): 1584–91.
CHAPTER 5
1. C. J. Lepre and D. V. Kent, ‘New Magnetostratigraphy for the Olduvai Subchron in the Koobi Fora Formation, Northwest Kenya, with Implications for Early Homo’, Earth and Planetary Science Letters 290 (2010): 362–74; I. McDougall et al., ‘New Single Crystal 40Ar/39Ar Ages Improve Time Scale for Deposition of the Omo Group, Omo-Turkana Basin, East Africa’, Journal of the Geological Society, London 169 (2011): 213–26.
2. The oldest known Asian Homo erectus are two from Mojokerto in Java and are thought to have lived around 1.81 ± 0.04 and 1.66 ± 0.04 myr. C. C. Swisher III et al., ‘Age of the Earliest Known Hominids in Java, Indonesia’, Science 263 (1994): 1118–21.
3. R. Dennell and W. Roebroeks, ‘An Asian Perspective on Early Human Dispersal from Africa’, Nature 438 (2005): 1099–1104.
4. See Humans Who Went Extinct, 53–4.
5. L. Gabunia et al., ‘Earliest Pleistocene Hominid Cranial Remains from Dmanisi, Republic of Georgia: Taxonomy, Geological Setting, and Age’, Science 288 (2000): 1019–25.
6. Finlayson, Neanderthals and Modern Humans.
7. Dennell and Roebroeks, ‘An Asian Perspective on Early Human Dispersal from Africa’.
8. D. Lordkipanidze et al., ‘Postcranial Evidence from Early Homo from Dmanisi, Georgia’, Nature 449 (2007): 305–10.
9. J. Hawks, ‘News flash: Dmanisi hominids were not short’,
10. J. Kappelman, ‘The Evolution of Body Mass and Relative Brain Size in Fossil Hominids’, Journal of Human Evolution 30 (1996): 243–76; F. Spoor et al., ‘Implications of New Early Homo Fossils from Ileret, East of Lake Turkana, Kenya’, Nature 448 (2007): 688–91.
11. Finlayson, The Humans Who Went Extinct.
12. L. C. Aiello and P. Wheeler, ‘The Expensive-Tissue Hypothesis’, Current Anthropology 36 (1995): 199–221.
13. C. V. Gisolfi and F. Mora, The Hot Brain: Survival, Temperature and the Human Body (Cambridge, Mass.: Bradford, 2000).
14. R. B. Eckhardt, ‘Was Plio-Pleistocene Hominid Brain Expansion a Pleiotropic Effect of Adaptation to Heat Stress?’, Anthropologischer Anzeiger 45 (1987): 193–201.
15. Known as the Oldowan.
16. Beyene, ‘Characteristics and Chronology’.
17. R. J. Blumenschine and C. R. Peters, ‘Archaeological Predictions for Hominid Land Use in the Paleo-Olduvai Basin, Tanzania, during Lowermost Bed II Times’, Journal of Human Evolution 34 (1998): 565–607; R. Potts et al., ‘Paleolandscape Variation and Early Pleistocene Hominid Activities: Members 1 and 7, Olorgesailie Formation, Kenya’, Journal of Human Evolution 37 (1999): 747–88; C. Shipton, ‘Taphonomy and Behaviour at the Acheulean Site of Kariandusi, Kenya’, African Archaeological Review 28 (2011): 141–55.
18. S. C. Antón, ‘Natural History of Homo erectus’, Yearbook of Physical Anthropology 46 (2003): 126–70.
19. J. C. A. Joordens et al., ‘Relevance of Aquatic Environments for Hominins: A Case Study from Trinil (Java, Indonesia)’, Journal of Human Evolution 57 (2009): 656–71.
20. Stegodon trigonocephalus.
21. E. A. Bettis III et al., ‘Way Out of Africa: Early Pleistocene Paleoenvironments Inhabited by Homo erectus in Sangiran, Java’, Journal of Human Evolution 56 (2009): 11–24.
22. Antón, ‘Natural History of Homo erectus’.
23. M. J. Morwood et al., ‘Fission-Track Ages of Stone Tools and Fossils on the East Indonesian Island of Flores’, Nature 392 (1998): 173–6; P. B. O’Sullivan et al., ‘Archaeological Implications of the Geology and Chronology of the Soa basin, Flores, Indonesia’, Geology 29 (2001): 607–10; A. Brumm et al., ‘Early Stone Technology on Flores and Its Implications for Homo floresiensis’, Nature 441 (2006): 624–8.
24. O. F. Huffman and Y. Zaim, ‘Mojokerto Delta, East Jawa: Paleoenvironment of Homo modjokertensis—First Results’, Journal of Mineral Technology 10 (2003): 1–32.
25. S. Pappu, ‘Early Pleistocene Presence of Acheulian Hominins in South India’, Science 331 (2011): 1596–9.
26. M. Belmaker and E. Tchernov, ‘New Evidence for Hominid Presence in the Lower Pleistocene of the Southern Levant’, Journal of Human Evolution 43 (2002): 43–56.
27. The dates for the site fall within the 1.95–1.77-myr bracket. M. Sahnouni et al., ‘Further Research at the Oldowan site of Ain Hanech, North-eastern Algeria’, Journal of Human Evolution 43 (2002): 925–37.
28. M. Sahnouni et al., ‘Ecological Background to Plio-Pleistocene Hominin Occupation in North Africa: The Vertebrate Faunas from Ain Boucherit, Ain Hanech and El-Kherba, and Paleosol Stable-Carbon-Isotope Studies from El-Kherba, Algeria’, Quaternary Science Reviews 30 (2011): 1303–17.
29. Fossil hominid and Oldowan technology. This is a cave site that is not informative about the ecological conditions outside the cave. Some mammals appear to have been butchered in the cave but the faunal list does not permit us to reconstruct the environment of these hominids. E. Carbonell et al., ‘The First Hominin of Europe’, Nature 452 (2008): 465–70.
30. O. Oms et al., ‘Early Human Occupation of Western Europe: Paleomagnetic Dates for Two Paleolithic Sites in Spain’, PNAS 97 (2000): 10666–70; D. Barsky et al., ‘Raw Material Discernment and Technological Aspects of the Barranco León and Fuente Nueva 3 Stone Assemblages (Orce southern Spain)’, Quaternary International 223–4 (2010): 201–19; I. Toro-Moyano et al., ‘The Oldest Human Fossil in Europe Dated to ca 1.4 Ma at Orce (Spain)’, Journal of Human Evolution (2013):
The Improbable Primate Page 16