.
31. G. R. Scott and L. Gibert, ‘The Oldest Hand-Axes in Europe’, Nature 461 (2009): 82–5.
32. J. Agustí et al., ‘The Early Pleistocene Small Vertebrate Succession from the Orce Region (Guadix-Baza Basin, SE Spain) and Its Bearing on the First Human Occupation of Europe’, Quaternary International 223–4 (2010): 162–9.
33. Gabunia et al., ‘Earliest Pleistocene Hominid Cranial Remains from Dmanisi’.
34. R. X. Zhu et al., ‘New Evidence on the Earliest Human Presence at High Northern Latitudes in Northeast Asia’, Nature 431 (2004): 559–62.
35. R. X. Zhu et al., ‘Early Evidence of the Genus Homo in East Asia’, Journal of Human Evolution 55 (2008): 1075–85.
36. Dennell and Roebroeks, ‘An Asian Perspective on Early Human Dispersal from Africa’.
CHAPTER 6
1. R. Dennell, The Palaeolithic Settlement of Asia (Cambridge: Cambridge University Press, 2009).
2. deMenocal, ‘African Climate Change’.
3. L. Barham and P. Mitchell (eds.), The First Africans: African Archaeology from the Earliest Toolmakers to Most Recent Foragers (Cambridge: Cambridge University Press, 2008).
4. Finlayson, Neanderthals and Modern Humans. Although flake technology akin to the Oldowan, which allowed a quick and flexible response to local variability of resources, meant that flakes were still being made into historical times in Africa, Barham and Mitchell (eds.), The First Africans.
5. Barham and Mitchell (eds.), The First Africans.
6. 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.
7. N. Goren-Inbar et al., ‘Pleistocene Milestones on the Out-of-Africa Corridor at Gesher Benot Ya’aqov, Israel’, Science 289 (2000): 944–7; N. Goren-Inbar et al., The Acheulian Site of Gesher Benot Ya’akov, Israel, i. The Wood Assemblage (Oxford: Oxbow Books, 2002).
8. Palaeoloxodon (Elephas) antiquus. N. Goren-Inbar et al., ‘A Butchered Elephant Skull and Associated Artifacts from the Acheulian Site of Gesher Benot Ya’akov, Israel’, Paléorient 20 (1994): 99–112.
9. Edible plants included wild grape Vitis sylvestris, water chestnut Trapa natans, prickly water lily Euryale ferox, cattail Typha spp., oak Quercus spp., wild pistachio Pistacia atlantica, wild olive Olea europaea, plum Prunus spp., and jujube Ziziphus spina-christi.
10. N. Goren-Inbar et al., ‘Nuts, Nut Cracking, and Pitted Stones at Gesher Benot Ya’aqov, Israel’, PNAS 99 (2002): 2455–60.
11. N. Goren-Inbar et al., ‘Evidence of Hominin Control of Fire at Gesher Benot Ya’aqov, Israel’, Science 304 (2004): 725–7.
12. R. Wrangham, Catching Fire: How Cooking Made Us Human (London: Profile Books, 2009).
13. Dennell, The Palaeolithic Settlement of Asia.
14. Two classic reviews by Clive Gamble summarize the European Middle Pleistocene: C. Gamble, The Palaeolithic Settlement of Europe (Cambridge: Cambridge University Press, 1986); C. Gamble, The Palaeolithic Societies of Europe (Cambridge: Cambridge University Press, 1999).
15. Typical European Middle Pleistocene large mammals associated with Homo erectus include deer, rhinos, elephants, bison, musk ox, reindeer, wolves, bears, lions, leopards, and hyenas.
16. S. A. Parfitt et al., ‘The Earliest Record of Human Activity in Northern Europe’, Nature 438 (2005): 1008–12.
17. C. Stringer, Homo britannicus: The Incredible Story of Human Life in Britain (London: Allen Lane, 2006).
18. Well summarized in Dennell, The Palaeolithic Settlement of Asia.
19. L. B. Vishnyatsky, ‘The Paleolithic of Central Asia’, Journal of World Prehistory 13 (1999): 69–122; A. Markova, ‘Fossil Rodents (Rodentia, Mammalia) from the Sel’Ungur Acheulean Cave Site (Kirghizstan)’, Acta Zoologica Cracoviensis 35 (1992): 217–39.
20. Summarized in Dennell, The Palaeolithic Settlement of Asia.
21. M. D. Petraglia, ‘The Lower Paleolithic of the Arabian Peninsula: Occupations, Adaptations, and Dispersals’, Journal of World Prehistory 17 (2003): 141–79.
22. L. A. Schepartz, ‘Upland Resources and the Early Palaeolithic Occupation of Southern China, Vietnam, Laos, Thailand and Burma’, World Archaeology 32 (2000): 1–13.
23. C. Finlayson et al., ‘Biogeography of Human Colonizations and Extinctions in the Pleistocene’, Memoirs Gibcemed 1 (2000): 1–69.
24. Finlayson, Neanderthals and Modern Humans.
25. We used the terms sources and sinks which were well established in metapopulation biology, where source populations are those that produce a surplus of individuals each breeding season. These individuals can disperse to other source populations and especially to sink populations. Sink populations are marginal populations that are only maintained by immigrants as the local conditions are insufficient to maintain a resident population.
26. N. A. Drake et al., ‘Ancient Watercourses and Biogeography of the Sahara Explain the Peopling of the Desert’, PNAS 108 (2011): 458–62.
27. C. Finlayson, Avian Survivors: The History and Biogeography of Palearctic Birds (London: T. & A. D. Poyser, 2011).
28. I discussed Homo heidelbergensis at length in The Humans Who Went Extinct so I limit the discussion here to the relevance of retaining Homo erectus as a more useful term. Finlayson, Avian Survivors and references therein.
29. A polytypic species is one that usually occupies a large geographical area and has significant regional variation. Populations at the extremes may appear quite different from each other but they are connected by others in between that interbreed and confirm that all populations, despite these differences, all belong to a single species.
CHAPTER 7
1. Finlayson, The Humans Who Went Extinct.
2. Barham and Mitchell (eds.), The First Africans. These authors provide a useful summary of the African evidence in the period between 430 and 70 kyr, which is the subject of this chapter.
3. Moments when population sizes declined severely, restricting their genetic diversity.
4. O. M. Pearson, ‘Postcranial Remains and the Origin of Modern Humans’, Evolutionary Anthropology 9 (2000): 229–47.
5. The Humans Who Went Extinct, 234 n. 46 lists references that confirm the endurance adaptation of the African erectus lineage.
6. At Omo Kibish in Ethiopia, dated to ~195 kyr. I. McDougall et al., ‘Stratigraphic Placement and Age of Modern Humans from Kibish, Ethiopia’, Nature 433 (2005): 733–6. But see also Preface.
7. Wolpoff, ‘The Systematics of Homo’.
8. We can consider the different lineages, based on anatomical differences, as subspecies of Homo sapiens. The subspecies is a taxonomic category that refers to populations of a single species that may be distinguished on the basis of differences of anatomy, colouration, voice, etc. They are different from each other but are able to interbreed and produce viable offspring. Using this convention, the African lineage would be Homo sapiens sapiens, the nominate subspecies. The Neanderthals, the Eurasian lineage, would be Homo sapiens neanderthalensis and erectus would be Homo sapiens erectus. Should we consider heidelbergensis to be sufficiently distinct from erectus to warrant subspecific status, then it would be Homo sapiens heidelbergensis. Similar criteria could be applied to other forms that are more contentious, e.g. ergaster, rhodesiensis, etc.
9. Generally referred to as Mode 3 technology, contrasting with Oldowan (Mode 1) and Acheulian (Mode 2). The previously prepared cores are referred to as Levallois cores. The technique allowed for greater control over the production of flakes; more could be extracted from a core, making the process more efficient than before. The method also permitted the making of stone-tipped spears, by hafting stone tools onto wooden shafts. These were the first composite tools known to have been made by Homo; Barham and Mitchell (eds.), The First Africans.
10. S. McBrearty, ‘Down with the Revolution’, in P. Mellars et al. (eds.), Rethinking the Human Revolution (Cambridge: McDonald Institute Monographs, 2007).
11. Robin Dunbar at the University of Liverpool has pioneered research into the importance of the brain in the context of the evolution of social groups. See, for example, R. Dunbar, The Human Story: A New History of Mankind’s Evolution (London: Faber and Faber, 2004) and discussion in The Humans Who Went Extinct, 207.
12. Drake et al., ‘Ancient Watercourses’; R. Tjallingii et al., ‘Coherent High- and Low-Latitude Control of the Northwest African Hydrological Balance’, Nature Geoscience 1 (2008): 670–5.
13. Finlayson, The Humans Who Went Extinct, 94–5.
14. Barham and Mitchell (eds.), The First Africans; V. Rots et al., ‘Aspects of Tool Production, Use, and Hafting in Palaeolithic Assemblages from Northeast Africa’, Journal of Human Evolution 60 (2011): 637–64.
15. Ammotragus lervia.
16. R. G. Klein and K. Scott, ‘Re-Analysis of Faunal Assemblages from the Haua Fteah and Other Late Quaternary Archaeological Sites in Cyrenaica, Libya’, Journal of Archaeological Science 13 (1986): 515–42.
17. A. Bouzouggar et al., ‘82,000-year-old Shell Beads from North Africa and Implications for the Origins of Modern Human Behaviour’, PNAS 104 (2007): 9964–9.
18. The authors found ochre and that the shells were perforated, suggesting modern human behaviour which was linked to the exploitation of the coast. I disagree with the idea of a sudden emergence of modern behaviour, a concept which is, in any case, slippery and impossible to define.
19. C. Marean et al., ‘Early Human Use of Marine Resources and Pigment in South Africa during the Middle Pleistocene’, Nature 449 (2007): 905–9.
20. R. G. Klein, The Human Career: Human Biological and Cultural Origins (Chicago: University of Chicago Press, 1999).
21. H. Faure et al., ‘The Coastal Oasis: Ice Age Springs on Emerged Continental Shelves’, Global and Planetary Change 33 (2002): 47–56.
22. J. E. Yellen et al., ‘The Archaeology of Aduma Middle Stone Age Sites in the Awash Valley, Ethiopia’, PaleoAnthropology 10 (2005): 25–100.
23. Barham and Mitchell (eds.), The First Africans.
CHAPTER 8
1. Estimates vary but most cluster around 400 kyr, see Humans Who Went Extinct, 106–7.
2. The argument that the Neanderthals originated in Europe from Middle Pleistocene ancestors is prevalent in the literature, e.g. C. Stringer and C. Gamble, In Search of the Neanderthals (London: Thames and Hudson, 1993). It stems from the number of Neanderthal fossils that have been recovered in Europe but this may be a simple artefact of the sampling, western Europe having been subjected to intense excavation for over a century while sites in other parts of the Neanderthal range, in Siberia for example, have been less intensely studied.
3. C. Finlayson and J. S. Carrión, ‘Rapid Ecological Turnover and Its Impact on Neanderthal and Other Human Populations’, Trends in Ecology and Evolution 22 (2007): 213–22.
4. C. Finlayson, ‘On the Importance of Coastal Areas in the Survival of Neanderthal Populations during the Late Pleistocene’, Quaternary Science Reviews 27 (2008): 2246–52.
5. C. Finlayson et al., ‘Late Survival of Neanderthals at the Southernmost Extreme of Europe’, Nature 443 (2006): 850–3. These last dates have been disputed on various grounds but have been robustly defended so that they remain the oldest known Neanderthal dates: J. Zilhao and P. Pettit, ‘On the New Dates for Gorham’s Cave and the Late Survival of Iberian Neanderthals’, Before Farming 3 (2006): 1–9; C. Finlayson et al., ‘Gorham’s Cave, Gibraltar—The Persistence of a Neanderthal Population’, Quaternary International 181 (2008): 64–71. Bradshaw and colleagues calculated the date when the Gorham’s Cave Neanderthals disappeared to 31.5 kyr. They based their estimate on the assumption that the last dates, around 32 kyr, could not represent the last ones but a small surviving population instead: C. Bradshaw et al., ‘Robust Estimates of Extinction Time in the Geological Record’, Quaternary Science Reviews 33 (2012): 14–19.
6. R. Pinhasi et al., ‘Revised Age of Late Neanderthal Occupation and the End of the Middle Paleolithic in the Northern Caucasus’, PNAS 108 (2011): 8611–16; R. E. Wood et al., ‘Radiocarbon Dating Casts Doubt on the Late Chronology of the Middle to Upper Palaeolithic Transition in Southern Iberia’, PNAS doi/10.1073/pnas.1207656110. These authors have redated Neanderthal sites across Europe using a new method of cleaning bone prior to radiocarbon dating. They have concluded that most resampled dates were older than under the previous untreated dates. From this they have concluded that the Neanderthals went extinct rapidly across Europe around 40 kyr. The conclusions have several flaws. In the first place these dates around 40 kyr must represent a healthy Neanderthal population if it was leaving so much evidence across the continent so the Neanderthals must have disappeared after 40 kyr. Second, their method of sampling requires that sufficient collagen is preserved to allow the material to be dated. Collagen survives best in cold and dry environments. This is precisely where we would expect the Neanderthals to have disappeared first. In contrast, warm and wet conditions, where the Neanderthals survived late, are unsuitable for this method of dating. So these results are biased towards the older dates. The late dates from Gorham’s Cave were taken from charcoal and not bone. When this team subjected the Gorham’s Cave charcoal to a similar cleaning treatment they found no difference in the treated and untreated dates: T. F. G. Higham et al., ‘The Radiocarbon Chronology of Gorham’s Cave’, in R. N. E. Barton, C. B. Stringer, and J. C. Finlayson (eds.), Neanderthals in Context: A Report of the 1995–1998 Excavations at Gorham’s and Vanguard Caves, Gibraltar (Oxford: Oxford University School of Archaeology Monograph 75, 2011).
7. T. Akazawa, K. Aochi, and O. Bar-Yosef, Neandertals and Modern Humans in Western Asia (New York: Plenum Press, 1998).
8. Finlayson and Carrión, ‘Rapid Ecological Turnover and Its Impact on Neanderthal and Other Human Populations’.
9. Hence why the most recent dates from these regions have not crossed this threshold, see n. 6.
10. The Spanish Region of Andalucía and the British Territory of the Rock of Gibraltar combined; R. Jennings et al., ‘Southern Iberia as a Refuge for the Last Neanderthal Populations’, Journal of Biogeography 38 (2011): 1873–85.
11. Using a Geographical Information System (GIS) we constructed two climate maps drawing on present-day temperature and rainfall measurements from 338 weather stations across the study area.
12. C. B. Stringer et al., ‘Neanderthal Exploitation of Marine Mammals in Gibraltar’, PNAS 105 (2008): 14319–24.
13. M. Cortés-Sánchez et al., ‘Earliest Known Use of Marine Resources by Neanderthals’, PLoS One 6 (2011): e24026.
14. A number of authors argue that brain-selective nutrients, particularly the omega-3 fatty acid docosahexaenoic acid DHA and arachidonic acid AA, have been essential in the successful development of the human brain. They extrapolate this ontogenetic aspect to phylogenetics inferring that the nutrients were necessary for the encephalization of the human brain; see S. C. Cunnane and K. M. Stewart (eds.), Human Brain Evolution: The Influence of Freshwater and Marine Food Resources (Hoboken, NJ, Wiley-Blackwell, 2010).
15. K. Milton, ‘Reply to S. C. Cunnane’, American Journal of Clinical Nutrition 72 (2000): 1586–7.
16. Among Neanderthal sites, 88 per cent had wetland birds, 28 per cent had coastal birds, and another 28 per cent had marine birds present. Many sites seem to have been important wetlands as the average number of wetland bird species represented was ~6. C. Finlayson et al., ‘The Homo Habitat Niche: Using the Avian Fossil Record to Depict Ecological Characteristics of Palaeolithic Eurasian Hominins’, Quaternary Science Reviews 30 (2011): 1525–32.
17. P. Mellars, ‘The Neanderthal Legacy: An Archaeological Perspective from Western Europe’ (Princeton: Princeton University Press, 1996);Jennings et al., ‘Southern Iberia’.
18. Finlayson, Neanderthals and Modern Humans.
19. Among Neanderthal sites 92 per cent incorporated rocky habitats with an average of ~11 rocky habitat bird species per site. Finlayson
et al., ‘The Homo Habitat Niche’.
20. F. J. Jiménez-Espejo et al., ‘Environmental Conditions and Geomorphologic Changes during the Middle-Upper Paleolithic in the Southern Iberian Peninsula’, Geomorphology 180–1 (2013): 205–16.
21. Studies of deep-sea marine cores, drilled from the seabed, provide useful information on climate. The different layers deposited in the seabed are picked up in the cores and can be dated. The chemical composition of the different layers informs about climatic conditions at the time of deposition.
The Improbable Primate Page 17