Sally McBrearty               The revolution that wasn’t: a new
Department of Anthropology,   interpretation of the origin...
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manipulation of symbols, and the origin of      two great Eur...
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 The earliest modern Europeans were                    ...
456                           .   . . 

fossil record and the genetic composition of    rather than b...
   ’                                   457

considered the temporal equivalent of the               ...
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a misapplication of a European model.             the genus ...
   ’                                459

record of the Middle Pleistocene (Howell,       works (Brau...
Table 1 Later African Hominidae, their archaeological associations and dates
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Site                            Specimen             Archaeology                   Dat...
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472



Table 1, Group 2    Continued

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Table 1   Continued

Site                         Specimen                Archaeology              Date            Method ...
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Table 1, Group 3       Continued

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Mcbrearty Brooks 00

  1. 1. Sally McBrearty The revolution that wasn’t: a new Department of Anthropology, interpretation of the origin of modern University of Connecticut, human behavior Storrs, Connecticut 06269, U.S.A. E-mail: Proponents of the model known as the ‘‘human revolution’’ claim mcbrearty@uconn.edu that modern human behaviors arose suddenly, and nearly simul- taneously, throughout the Old World ca. 40–50 ka. This fundamental Alison S. Brooks behavioral shift is purported to signal a cognitive advance, a possible Department of Anthropology, reorganization of the brain, and the origin of language. Because the George Washington earliest modern human fossils, Homo sapiens sensu stricto, are found in University, Washington, Africa and the adjacent region of the Levant at >100 ka, the ‘‘human DC 20052, U.S.A. E-mail: revolution’’ model creates a time lag between the appearance of abrooks@gwu.edu anatomical modernity and perceived behavioral modernity, and creates the impression that the earliest modern Africans were behav- Received 3 June 1999 iorally primitive. This view of events stems from a profound Euro- Revision received 16 June centric bias and a failure to appreciate the depth and breadth of the 2000 and accepted 26 July African archaeological record. In fact, many of the components of 2000 the ‘‘human revolution’’ claimed to appear at 40–50 ka are found in the African Middle Stone Age tens of thousands of years earlier. Keywords: Origin of Homo These features include blade and microlithic technology, bone tools, sapiens, modern behavior, increased geographic range, specialized hunting, the use of aquatic Middle Stone Age, African resources, long distance trade, systematic processing and use of archaeology, Middle pigment, and art and decoration. These items do not occur suddenly Pleistocene. together as predicted by the ‘‘human revolution’’ model, but at sites that are widely separated in space and time. This suggests a gradual assembling of the package of modern human behaviors in Africa, and its later export to other regions of the Old World. The African Middle and early Late Pleistocene hominid fossil record is fairly continuous and in it can be recognized a number of probably distinct species that provide plausible ancestors for H. sapiens. The appearance of Middle Stone Age technology and the first signs of modern behavior coincide with the appearance of fossils that have been attributed to H. helmei, suggesting the behavior of H. helmei is distinct from that of earlier hominid species and quite similar to that of modern people. If on anatomical and behavioral grounds H. helmei is sunk into H. sapiens, the origin of our species is linked with the appearance of Middle Stone Age technology at 250–300 ka. 2000 Academic Press Journal of Human Evolution (2000) 39, 453–563 doi:10.1006/jhev.2000.0435 Available online at http://www.idealibrary.com on Introduction and background 2000; Diamond, 1992; Mellars, 1995, 1996; Nobel Davidson, 1991; Tattersall, The human revolution in Europe 1995; Bar-Yosef, 1998). The ‘‘human For at least the past 15 years, most recon- revolution’’ model proposes a dramatic structions of later human evolutionary alteration in human behavior at the Middle history have featured a relatively brief and Paleolithic to Upper Paleolithic transition at dramatic shift known as the ‘‘human revol- about 40 ka. This behavioral breakthrough ution’’ (Binford, 1985, 1989; Mellars is thought by some to correspond to Stringer, 1989; Klein, 1989a, 1994, 1995, increased cognitive sophistication, the 0047–2484/00/110453+111$35.00/0 2000 Academic Press
  2. 2. 454 .   . .  manipulation of symbols, and the origin of two great European wars, and the trend language (e.g., White, 1982; Mellars in archaeology in the second half of the Stringer, 1989; Diamond, 1992; Byers, twentieth century has been the study of 1994; Mithen, 1994, 1996; Klein, 1995; but local sequences and the application of see Kay et al., 1998). We believe that the models of cultural evolution (Otte Keeley, model of the ‘‘human revolution’’ is fatally 1990). flawed. Modern humans and modern In terms of developments in world pre- human behaviors arose first in Africa, and history, however, Western Europe is a we examine the African record to reveal a remote cul de sac with a somewhat anom- different picture of the nature of events. alous prehistoric record. We argue here that The concept of a ‘‘human revolution’’ and models derived from the unique record of the periodization of Stone Age prehistory European prehistory do not explain events have their roots in the nineteeth-century in Africa where the origin of modern people probings of the Western European archaeo- actually occurred. In the Holocene, western logical record. The first paleolithic classifi- Europe experienced a series of incursions catory schemes were based on the Western from the less peripheral portions of the Old European large mammal succession (Lartet World. Each arrival of a wave of invaders Christy, 1865–1875; Lyell, 1868), and and alien technology induced a fairly these authors emphasized the wide techno- sudden, rapid cultural turnover. These logical gulf separating the l’age du renne disruptive episodes are reflected in the (Upper Paleolithic) from the earlier phases European archaeological record as dis- (Lartet Christy, 1865–1875:25). By the continuities that punctuate industrial 1920s the concept of an Upper Paleolithic periods of relatively long duration. They distinguished by the appearance of engrav- have been sometimes described as ‘‘revol- ing, sculpture, painting, beads, and worked utions,’’ such as the ‘‘neolithic revolution’’ bone tools had become current. A tripartite of Childe (1936, 1942). division into Lower, Middle and Upper Recent paleoclimatic data and refined Paleolithic based upon stone tool technology chronologies have supported the early sug- (De Mortillet, 1900; Obermeier, 1924; gestion of Howell (1951) that regions of Burkitt, 1921, 1928, 1933; Kendrick, 1925; Pleistocene Europe were repeatedly isolated Menghin, 1931) echoed the three-age sys- by ice and mountain barriers, so that its tems of Thomsen (1837) and Worsaae hominid populations were periodically (1849) that partitioned the total prehistoric reduced or even eliminated (Howell, 1952; record into ages of Stone, Bronze, and Iron. Gamble, 1986, 1994; Jochim, 1987; The Lower, Middle and Upper Paleolithic Hublin, 1998a). Moreover, it has become divisions of the western European record increasingly clear that the Neanderthals have continued to dominate discourse in the were replaced by modern humans in Europe field, despite problems in the application of within too short a period for the former to these divisions to sequences in Eastern and have evolved into the latter (Mellars, Southern Europe (Morselli, 1926:292). 1998a,b, 1999, Bocquet-Appel Demars, None of these temporal divisions was 2000). Thus, the ‘‘revolutionary’’ nature of intended as an evolutionary scheme, but the European Upper Paleolithic is most rather they were thought to reflect repeated probably due to discontinuity in the invasions by outsiders with new ideas. archaeological record rather than to the sort Perhaps not surprisingly, a picture of of rapid cultural, cognitive, and/or biological Europe conquered by invaders with superior transformation that has been argued by technology had little appeal in the light of proponents of the ‘‘human revolution.’’
  3. 3.    ’ 455 The earliest modern Europeans were The fossil evidence for an African origin Africans for modern humans is robust. It is clear that modern humans (H. sapiens sensu stricto) Who were the earliest modern Europeans? It were certainly present in Africa by 130 ka is becoming increasingly difficult to deny (Day Stringer, 1982; Deacon, 1989), and that they were Africans. Although the perhaps as early as 190 ka if specimens ‘‘mitochondrial Eve’’ hypothesis, first such as Singa are considered modern articulated by Cann et al. (1987), has been (McDermott et al., 1996; Stringer, 1996). revised in light of criticism (Templeton, Modern humans do not appear in Europe or 1992; Hedges et al., 1992; Ayala, 1995), and Central Asia before ca. 40 ka; earliest dates population size and structure have effects on for the Levant range between ca. 80 ka and the distribution of genetic characters that 120 ka (Day, 1969, 1972; Day Springer, were not taken into account in early recon- 1982, 1991; Stringer, 1989, 1992; structions (Harpending et al., 1993, 1998; McBrearty, 1990b; Stringer et al., 1989; Sherry et al., 1994; Relethford, 1995; Brauer, 1984a,b, 1989; Stringer Andrews, ¨ Relethford Harpending, 1995), genetic 1988; Valladas et al., 1988; Grun ¨ data either directly support or are consistent Stringer, 1991; Miller et al., 1991; Foley with an African origin for modern humans Lahr, 1992; Mercier et al., 1993; Deacon, (Wainscoat et al., 1986; Cann, 1988; 1993b; Brooks et al., 1993a,b; Stringer, Stringer Andrews, 1988; Vigilant et al., 1993a; Schwarcz, 1994; Straus, 1994; 1991; Stoneking, 1993; Stoneking et al., Bar-Yosef, 1994, 1995a, 1998; but see 1993; Relethford Harpending, 1994; Howells, 1989). Recent evidence suggests Ayala, 1995; Nei, 1995; Goldstein, 1995; that modern humans were present in Tishkoff et al., 1996; Ruvolo, 1996, 1997; Australia as early as 62 ka (Stringer, 1999; Irish, 1998; Pfeiffer, 1998; Zietkiweicz et al., Thorne et al., 1999). 1997; Pritchard et al., 1999; Quintana- Although some, notably Brauer (1984a,b, ¨ Murci, 1999; Relethford Jorde, 1999; 1989), favor a scenario involving some inter- Tishkoff et al., 2000; see Relethford, 1998 breeding among Neanderthal and modern and Jorde et al., 1998 for recent reviews). human populations, the successful extrac- As Howell (1994:306) observes, ‘‘The tion and analysis of fragmentary mito- phylogenetic roots of modern humans are chondrial DNA (mtDNA) from both the demonstrably in the Middle Pleistocene. Neanderthal type fossil (Krings et al., 1997, The distribution of those antecedent 1999) and additional material from the populations appear to lie outside of western northern Caucasus (Ovchinnikov et al., and eastern Eurasia, and more probably 2000) appears to remove the Neanderthals centered broadly on Africa.’’1 from modern human ancestry. Body propor- tions of early European H. sapiens fossils 1. The Middle to Late Pleistocene boundary is the suggest a tropical adaptation and support an beginning of the last interglacial, at approximately African origin (Holliday Trinkaus, 1991; 130 ka; the base of the Middle Pleistocene is the shift from reversed to normal magnetic polarity at the Ruff, 1994; Pearson, 1997, 2000; Holliday, Matuyama–Brunhes boundary, dated to about 780 ka 1997, 1998, 2000). A single migration or (Butzer Isaac, 1975; Imbrie Imbrie, 1980; Berger population bottleneck was originally envis- et al., 1984; Martinson et al., 1987; Shackleton et al., 1990; Deino Potts, 1990; Cande Kent, 1992; aged in the ‘‘African Eve hypothesis’’ (Cann Baksi et al., 1992; Tauxe et al., 1992). Further evidence et al., 1987), but a succession of population may confirm recent suggestions (Schneider et al., 1992; dispersals, subsequent isolation induced by Singer Pringle, 1996; Hou et al., 2000) that the age of this geomagnetic polarity reversal be revised to climatic events and local adaptation may ca. 790 ka. better account for the complexity of the
  4. 4. 456 .   . .  fossil record and the genetic composition of rather than by genetic processes, the most present human populations (Howells, 1976, likely scenario would be an accretionary 1989, 1993; Boaz et al., 1982; Foley Lahr, process, a gradual accumulation of modern 1992; Lahr Foley, 1994, 1998; Ambrose, behaviors in the African archaeological 1998b). record (cf. Allsworth-Jones, 1993). This It can be deduced from the archaeological change need not be unidirectional or con- evidence that on a continent-wide scale the fined to a single location. Rather, we might African record differs markedly from that expect innovative behaviors to appear at of Europe in its degree of population con- different times and in different regions, and tinuity. While parts of Africa, such as the due to low population densities we might Sahara or the interior of the Cape Province expect the transmission of new ideas to be of South Africa, do appear to have experi- sporadic. enced interruptions in human settle- As early as the 1920s it was clear that the ment during glacial maxima (Deacon African archaeological record could not be Thackeray, 1984; Williams, 1984; Butzer, accommodated within the European Paleo- 1988b; Brooks Robertshaw, 1990; lithic model. A separate scheme of Earlier, Mitchell, 1990), climatic reconstructions Middle and Later Stone Ages (ESA, MSA, suggest that the contiguous expanse of and LSA) was devised for Stone Age Africa steppe, savanna and woodland biomes avail- (Goodwin van Riet Lowe, 1929) to able for human occupation, especially in the emphasize its distinctiveness from the tropical regions of the continent, was always Lower, Middle, and Upper Paleolithic of substantially larger than the comparable Europe. The ESA, MSA and LSA were first regions in Europe. Perhaps as a result, defined on technological grounds on the hominid populations in Africa, while prob- basis of material from South Africa ably widely dispersed, appear to have (Goodwin, 1928; Goodwin van Riet been consistently larger (Relethford Lowe, 1929). The terms were formally Harpending, 1995; Jorde et al., 1998; endorsed by the Panafrican Congress of Relethford Jorde, 1999; Tishkoff et al., 1955 (Clark, 1957a: xxxiii). The ESA as it is 2000). now understood includes both the Oldowan and the Acheulian; the MSA encompasses flake and blade tool industries which often Revolution or evolution? The African include prepared cores and points; and the data LSA is characterized by microlithic tech- How might the archaeological signature of nology. The MSA was distinguished by the continuous evolutionary change be expected presence of prepared core technology and, to differ from that of abrupt replacement? If at most sites, unifacial and/or bifacial projec- the entire human species experienced a sim- tile points, and by the absence of handaxes ultaneous, punctuated, genetically encoded and microliths, hallmarks of the Acheulian event, such as the development of modern and LSA respectively. capacities for language (Klein, 1995; Before 1972, in the absence of accurate Diamond, 1992), one would expect the chronometric dates, a radiocarbon date of transition to modern human behavior to be 60 ka from Acheulian levels at Kalambo abrupt, in Africa as well as in Europe and Falls, Zambia (Clark, 1969) was not Asia. On the other hand, if aspects of recognized as infinite. This frequently cited modern human culture in Africa were devel- date was particularly influential in establish- oped by hominids using existing cognitive ing the impression of a short chronology capabilities and transmitted by cultural for Africa. The MSA, at 60 ka, was
  5. 5.    ’ 457 considered the temporal equivalent of the quently retired (Bishop Clark, 1967: Upper Paleolithic of Europe. Therefore the 987), when a mixture of different occu- discovery of anatomically modern human pation levels was found to have occurred remains associated with MSA artefacts at during excavation at the ‘‘Second Inter- the South African sites of Border Cave mediate’’ type site of Magosi (Wayland and Klasies River occasioned no surprise.2 Burkitt, 1932; Clark, 1957b; Hole, 1959; The degree of regional differentiation, the Cole, 1967). Yet anachronisms, as well as ubiquitous presence of blades and blade long periods of transition between stages, cores, and the sophistication of projectile remain as problems (Vishnyatsky, 1994). point technology in the African MSA were A fairly abrupt MSA–LSA transition is considered comparable to the European apparent in the Mediterranean zones at the Upper Paleolithic. However, bone tools, art northern and southern margins of Africa. objects and beads were sparse when com- This seems consistent with the significant pared to the European Upper Paleolithic, documented gaps in the settlement history particularly the late Upper Paleolithic. of both regions (Close et al., 1990; Wendorf The rarity of elements regarded as critical et al., 1990, 1993a; Mitchell, 1990; Deacon to modern human culture in the MSA Thackeray, 1984; Klein, 1989b: 307). served as grounds for regarding Africa as a However, at rock shelter sites in tropical ‘‘cultural backwater,’’ the place that initially Africa with relatively continuous occu- gave rise to humanity, but failed to nurture pational records, such as Mumba, Tanzania its later development (e.g., Butzer, 1971; (Mehlman, 1979, 1989), Matupi, D. R. cf. Clark, 1975). In the later 1970s, new Congo (van Noten, 1977) and White dating techniques and more accurate Paintings, Botswana (Robbins Murphy, climatic correlations pushed back the age of 1998; Robbins et al., under review) there is a the MSA well beyond 100 ka. The MSA was gradual transition from MSA to LSA tech- recognized as the temporal equivalent of the nology over as much as 30 ka. Mehlman European Middle Paleolithic, not the Upper (1991) has urged the development of new Paleolithic. Attention focused on the human paradigms to accommodate the lack of a fossils associated with the MSA, which were punctuated event. now thought to be anomalously modern Because of the late, sudden, and nearly in appearance. The fact that many MSA simultaneous appearance in Europe of mod- artefacts recalled the Upper Paleolithic of ern humans and complex behavior, archae- Europe in both form and technology was ologists working in Africa have sought a forgotten. similar ‘‘human revolution’’ there. The fully The use of a classifactory scheme developed signature of modern human designed for Africa did not entirely remove behavior, including planning, sophisticated ambiguity, as many industries displayed technology and resource use, and symbolic characteristics of two different stages and behavior in the form of decorative art is could not be assigned to one of the three clearly present in the African LSA. As a re- divisions. Long transitional periods or sult, the MSA–LSA transition has been con- ‘‘Intermediates’’ were added to the tripartite flated with the Middle to Upper Paleolithic ESA–MSA–LSA scheme at the 1955 and the emergence of modern human behav- Panafrican Congress (Clark, 1957a: xxxiii), ior. Consequently the earliest anatomically but the ‘‘Intermediate’’ concept was subse- modern humans, which occur in MSA con- texts, are not accepted as fully ‘‘human’’. 2. The name Klasies River rather than Klasies River Mouth or KRM is adopted here to conform with the We suggest that the expectation of a recent usage of Hilary Deacon and his team. ‘‘human revolution’’ in Africa is ultimately
  6. 6. 458 .   . .  a misapplication of a European model. the genus Homo when compared to the more Further, we reject the idea of a time lag derived state in the Neanderthals (Rak, between anatomical and behavioral change 1993). in Africa, such as that proposed by Klein Specimens formerly attributed to (1992, 1994, 1995, 1998). There was no ‘‘archaic’’ H. sapiens exhibit a number of ‘‘human revolution’’ in Africa. Rather, in plesiomorphic traits, including long low this paper we present data from the human crania, large brow ridges, large, prognathic fossil and archaeological records to show faces with large teeth, and the lack of a chin. that novel features accrued stepwise. The chief justification for the inclusion of Distinct elements of the social, economic, these fossils in our species has been their and subsistence bases changed at different large brain sizes, though brain size is in part rates and appeared at different times and a function of body mass, known to be quite places. We describe evidence from the large among these hominids (Grine et al., African MSA to support the contention that 1995; Ruff et al., 1997; Kappelman, 1997). both human anatomy and human behavior Recent discussions of later hominid phy- were intermittently transformed from an logeny (e.g., Stringer, 1993b, 1994, 1995, archaic to a more modern pattern over a 1996; Lahr Foley, 1994; Foley Lahr, period of more than 200,000 years. 1997; Rightmire, 1998) have recognized the distinctiveness of non-Neanderthal Middle Pleistocene hominids and have resurrected The hominid fossil record the taxon H. heidelbergensis Schoetensack, Until recently, most reconstructions of later 1908 for them, but we question the human phylogeny recognized only one attribution of the African material to this species after H. erectus. Grade-based taxon. schemes commonly divided H. sapiens into Paradoxically, H. sapiens Linnaeus, 1758 two variants, ‘‘archaic’’ H. sapiens and ‘‘ana- lacks a satisfactory definition. Howell (1978: tomically modern’’ H. sapiens (H. sapiens 201) observed over 20 years ago, sensu stricto). The Neanderthals were then ‘‘The extensive relevant literature reveals an sometimes distinguished from other unexpected lack of concern with the bio- ‘‘archaic’’ H. sapiens at the subspecific logical distinctiveness of a now-dominant level as H. sapiens neanderthalensis (e.g., mammalian species’’, Campbell, 1964). We concur with such and the situation is virtually unchanged authors as Tattersall (1986, 1992), Kimbel today. The anatomy of H. sapiens is charac- (1991), Harrison (1993), Rak (1993) and terized by a high round cranium, a chin, a Stringer (1994, 1996) that there are grounds small orthognathic face, as well as reduced for distinguishing ‘‘archaic’’ from ‘‘modern’’ masticatory apparatus and brow ridges. It H. sapiens at the species level, and thus has been argued that most of these features we regard the appearance of ‘‘modern’’ H. can be explained by greater flexion in the sapiens as a speciation event. Here, we treat basicranium of H. sapiens (Lieberman, the Neanderthals as the distinct species H. 1998b; Spoor et al., 1999). neandertalensis King, 1864, and use the Because early fossils of H. sapiens dating name H. sapiens to refer only to H. sapiens to 130 ka, and perhaps as early as 190 ka, senus stricto. The use of the ‘‘anatomically are found in Africa (Grun et al., 1990; ¨ modern’’ label for H. sapiens sensu stricto is Deacon, 1989, 1993b; Day Stringer, not only unnecessary but also misleading, as 1982; McDermott et al., 1996), it is reason- many of the cranial features used to dis- able to seek evidence for the processes lead- tinguish H. sapiens are in fact primitive for ing to the origin of H. sapiens in the African
  7. 7.    ’ 459 record of the Middle Pleistocene (Howell, works (Brauer, 1984a,b, 1989; Smith, 1985, ¨ 1994). Although often described as 1993; Clark, 1988; Klein, 1989b, 1994; ‘‘scrappy’’ or insubstantial, the African Stringer, 1993a). Data in Table 1 roughly hominid fossil sample from this time period follow Day’s tripartite construct, though this numbers several dozen individuals (Table 1, should not be construed as an endorsement Figure 1). While the circumstances of for anagenesis or a grade-based taxonomy. recovery for some of the specimens are far Both the ascription of fossils to group and from ideal, this is unfortunately true for the attachment of taxonomic labels are many fossil discoveries, and in fact a fair problematic, and Group 1 specimens number of the African specimens were probably belong to several different species recovered by controlled excavation (e.g., (e.g., H. louisleakeyi, H. rhodesiensis). Ndutu, Cave of Hearths, Haua Fteah, The principal unresolved issue in the Mumba, Ngaloba, Klasies, Kapthurin post- clarification of the evolutionary relationships cranials). For others, stratigraphic context of the hominids in Group 1 is the enigmatic can be reasonably inferred, despite the fact status of H. erectus. This species is believed that they are surface finds (e.g., Kapthurin by many to have been confined to Asia mandibles, Eyasi, Buia). (Andrews, 1984; Tattersall, 1986; Groves, Howell (1994: 305f) has deemed the sol- 1989; Clarke, 1990; Kimbel, 1991; Larick ution of the evolutionary relationships Ciochon, 1996). Following Wood (1991, among later Middle Pleistocene hominid 1992), some now ascribe to H. ergaster populations one of the central problems in African fossils in the 1·5–2 Ma age range the study of human evolution, and the taxo- formerly attributed to H. erectus. Other nomic status of the African fossils is much authors (e.g., Rightmire, 1990, 1994, 1995, debated (e.g., Tattersall, 1986; Clarke, 1998; Brauer Mbua, 1992; Harrison, ¨ 1990; Foley, 1991a; Kimbel, 1991; 1993; Walker, 1993; Brauer, 1994) regard ¨ Stringer, 1992, 1993a, 1994, 1996; Aiello, H. erectus as a single polytypic species dis- 1993; Foley Lahr, 1997; Lahr Foley, tributed throughout most of the Old World, 1998; Rightmire, 1998). Revision of fossils and African specimens in our Group 1, ascribed to Homo (Wood, 1991, 1992; spanning a broad range of time, continue Wood Collard, 1999) has resulted in a to be ascribed to this taxon (e.g., OH9, more ‘‘bushy’’ or speciose taxonomic Kapthurin, in Wood, 1992). Attribution of picture for our genus in the Pliocene and the African and Asian Middle Pleistocene Early Pleistocene, but for the African material to a single species assumes an Middle and Late Pleistocene a unilineal adequate degree of gene flow to prevent model is often invoked. It is our belief that speciation, but the archaeological differ- the number of African Middle and Late ences between the regions suggest long term Pleistocene hominid species has been under- isolation (Schick, 1994; but see Hou et al., estimated, because behavioral and repro- 2000). ductive isolation may precede changes in the Group 1 in our scheme includes the bony skeleton (Tattersall, 1986, 1992, Kabwe (Broken Hill) cranium, type speci- 1993; Rak, 1993). men of H. rhodesiensis Woodward, 1921, as Over 25 years ago, Day (1973) suggested well as OH9. The latter specimen is usually separating African Middle and Late referred in the literature to H. erectus, but Pleistocene hominids into ‘‘early,’’ ‘‘inter- Louis Leakey (1961, 1963) emphatically mediate,’’ and ‘‘modern’’ groups, and this rejected this position. He saw the origin of grade-based practice has been followed, H. sapiens as a strictly African phenom- explicitly and implicitly, in many subsequent enon, and regarded OH9 as morphologically
  8. 8. Table 1 Later African Hominidae, their archaeological associations and dates 460 Site Specimen Archaeology Date Method Selected references Group 1 (H. erectus, H. ergaster, H. louisleakeyi, H. rhodesiensis) Aın Maarouf ¨ Left femoral shaft Acheulian Early Middle Associated fauna Hublin, 1992; (El Hajeb), Morocco Pleistocene Geraads et al., 1992 Berg Aukas, Namibia Femoral fragment None Undated Grine et al., 1995 Bodo, Ethiopia Adult cranium, Acheulian Mid to later Middle Associated fauna Conroy et al., 1978; parietal, distal humerus Pleistocene, 350 ka Kalb et al., 1980, 1982a,b; Asfaw, 1983; Clark et al., 1984; Rightmire, 1996 40 cf. Oldowan 640 ka–550 ka Ar/39Ar, associated Clark et al., 1994 fauna Buia, Danakil (Afar) Adult cranium, 2 None reported 1·0 Ma Paleomagnetism, Abbate et al., 1998 Depression, Eritrea incisors, pelvic associated fauna fragments Cave of Hearths, Mandible, radius Acheulian Early Late Associated fauna Cooke, 1962; Mason, 1962; .   . .  South Africa Pleistocene Mason et al., 1988; Tobias, 1971; Partridge, 1982; Pearson Grine, 1997 End Middle Associated fauna Howell, 1978 Pleistocene Eyasi, Tanzania Cranial fragments Sangoan 130 ka Extrapolation from Cooke, 1963; Mehlman, representing 3–4 overlying 14C dates, 1984, 1987 individuals underlying 230Th/234U dates, faunal correlation
  9. 9. Table 1, Group 1 Continued Site Specimen Archaeology Date Method Selected references Kabwe (Broken Hill), Adult cranium (E686), ?Sangoan or MSA? 125 ka Associated fauna Woodward, 1921; Pycraft Zambia (holotype of cranial, maxillary et al., 1928; Oakley, 1957; H. rhodesiensis) dental and postcranial Clark, 1959; Clark et al., (humeral, pelvic, 1950, 1968; Klein, 1973, femoral, tibial) remains 1994; Santa Luca, 1978; of d3 individuals Partridge, 1982; Vrba, 1982; Stringer, 1986 110 ka Aspartic acid Bada et al., 1974 racemization on hominid femoral fragment EM 793 700–400 ka Associated fauna (cf. Klein, 1994; Rightmire, Olduvai Beds III–IV) 1998 780 ka–1·33 Ma Associated fauna (cf. This paper, based upon Olduvai Beds III–IV). Klein, 1973, 1994; Correlation of top of Partridge, 1982; Hay, 1976; Bed IV with Matuyama Walter et al., 1991, 1992;    ’ Brunhes boundary Tamrat et al., 1995; Kimbel, 1995; Delson van Couvering, 2000 1·07–1·33 Ma Associated fauna (cf. This paper, based upon Olduvai Beds III–IV). Klein, 1973, 1994; Correlation of normal Partridge, 1982; Hay, 1976; polarity paleomagnetic Walter et al., 1991, 1992; zone at base of Masek Tamrat et al., 1995; Beds with Jaramillo Kimbel, 1995; Delson subchron van Couvering, 2000 461
  10. 10. Table 1, Group 1 Continued 462 Site Specimen Archaeology Date Method Selected references Kapthurin (Baringo) Two adult mandibles, Undiagnostic 230–780 ka K/Ar, associated fauna, Leakey et al., 1969; van Kenya (KNM-BK 67, 8518) paleomagnetism Noten, 1982; Howell, 1982; postcranials (ulna, van Noten Wood, 1985; talus, manus phalanges, Wood van Noten, 1986; KNM-BK 63–66) Tallon, 1978; Dagley et al., 1978; Rightmire, 1980; Solan Day, 1992; Wood, 1992; Groves, 1998 500–550 ka Deino McBrearty, under review 230 ´ Kebibat (Rabat), Subadult calvaria, None 200 ka Th/234U Stearns Thurber, 1965 Morocco maxillary fragment, mandible 300 ka–1·0 Ma Associated fauna Saban, 1975, 1977; Howell, 1978; Sausse, 1975b 40 Lainyamok, Kenya Femoral shaft, isolated Undiagnostic 390–330 ka Ar/39Ar Shipman et al., 1983; Potts teeth et al., 1988; Potts Deino, 1995 .   . .  Loyangalani Maxillary and None Late Middle/Early Associated fauna Twiesselmann, 1991 mandibular dentition Late Pleistocene ´ Melka Konture, Cranial fragments Acheulian Middle Pleistocene Associated fauna Chavaillon et al., 1974; Ethiopia Howell, 1978; Chavaillon, 1982 Ndutu, Tanzania Adult cranium cf. Acheulian 500–600 ka AAR on associated Mturi, 1976; Rightmire, mammalian bone 1980, 1983; Clarke, 1976, 1990
  11. 11. Table 1, Group 1 Continued Site Specimen Archaeology Date Method Selected references Ndutu, Tanzania 490–780 ka Upper limit: correlation This paper, based upon continued of reversed polarity Tamrat et al., 1995 excursion in underlying Norkilili Member of Masek Beds with Emperor subchron. Lower limit: correlation of top of Bed IV with Matuyama Brunhes boundary 370–990 ka Upper limit: 40Ar/39Ar This paper, based upon on Kerimasi caldera, Hay, 1976, 1990; Leakey probable source of et al., 1972; Leakey Hay, Norkilili Member of 1982; Clarke, 1976, 1990; Masek Beds. Lower Manega, 1993; Tamrat limit: correlation of et al., 1995; Walter et al., normal polarity 1991, 1992; Kimbel, 1995 paleomagnetic zone at base of Masek Beds with Jaramillo subchron    ’ OH 9, LLK, Olduvai Adult partial cranium Developed Oldowan or 1·15 Ma (surface, top K/Ar, paleomagnetism Hay, 1963, 1973, 1976, Gorge, Tanzania Acheulian of Bed II) 1990; Leakey, 1961, 1963; (type of H. Leakey, 1971a,b; Leakey louisleakeyi) Hay, 1982; Rightmire, 1979a, 1980, 1990, 1994; Wood, 1994 0·7–1·0 Ma K/Ar, paleomagnetism Hay, 1971 40 1·33–1·48 Ma Ar/39Ar, Hay, 1976, 1990; Manega, paleomagnetism 1993; Tamrat et al., 1995; Walter et al., 1991, 1992; Kimbel, 1995; White, 2000 463
  12. 12. Table 1, Group 1 Continued 464 Site Specimen Archaeology Date Method Selected references OH 11, DK, Olduvai Palate, maxilla None; both ca. 400 ka (surface, K/Ar, paleomagnetism Rightmire, 1979a; Leakey Gorge, Tanzania Acheulian MSA probably lower Ndutu inferred sedimentation Hay, 1982; Hay, 1994 elsewhere in Ndutu Beds) rates Beds ca. 490 ka Paleomagnetism; This paper, based upon correlation of reversed Tamrat et al., 1995 polarity excursion in underlying Norkilili Member of Masek Beds with Emperor subchron. 40 ca. 370 ka Ar/39Ar on Kerimasi Hay, 1976, 1990; Manega, caldera, probable 1993; Tamrat et al., 1995; source of Norkilili Walter et al., 1991, 1992; Member of Masek Kimbel, 1995; White, 2000; Beds. Delson van Couvering, 2000 OH 12, VEK, Olduvai Palate, maxilla, cranial None; Acheulian 780–620 ka (upper Bed K/Ar, paleomagnetism, Rightmire, 1979a; Leakey Gorge, Tanzania fragments elsewhere in Bed IV IV) inferred sedimentation Hay, 1982; Hay, 1994; rates Leakey Roe, 1994 .   . .  780 ka–1·2 Ma Paleomagnetism; This paper, based upon correlation of top of Tamrat et al., 1995 Bed IV with Matuyama Brunhes boundary 40 1·07–1·2 Ma Ar/39Ar Hay, 1976, 1990; Manega, paleomagnetism; 1993; Tamrat et al., 1995; correlation of normal Walter et al., 1991, 1992; polarity paleomagnetic Kimbel, 1995; White, 2000; zone at base of Masek Delson van Couvering, Beds with Jaramillo 2000 subchron
  13. 13. Table 1, Group 1 Continued Site Specimen Archaeology Date Method Selected references OH 22, VEK/MNK, Partial mandible None; Acheulian 800–600 ka (surface, K/Ar, paleomagnetism, Day, 1986; Leakey Hay, Olduvai Gorge, elsewhere in Bed IV Bed IV) inferred sedimentation 1982; Leakey Roe, 1994; Tanzania and overlying Ndutu rates Rightmire, 1979a, 1990; Beds Hay, 1994 780 ka–1·2 Ma Paleomagnetism; This paper, based upon correlation of top of Tamrat et al., 1995 Bed IV with Matuyama Brunhes boundary 40 1·07–1·2 Ma Ar/39Ar, Hay, 1976, 1990; Manega, paleomagnetism; 1993; Tamrat et al., 1995; correlation of normal Walter et al., 1991, 1992; polarity paleomagnetic Kimbel, 1995; White, 2000; zone at base of Masek Delson van Couvering, Beds with Jaramillo 2000 subchron OH 23, FLK, Olduvai Mandibular fragment Acheulian 400–600 ka (In situ, K/Ar, paleomagnetism, Day, 1986; Rightmire, Gorge, Tanzania Masek Beds) inferred sedimentation 1990; Leakey Roe, 1994; rates Hay, 1994    ’ 490–780 ka Paleomagnetism. This paper, based upon Upper limit: correlation Tamrat et al., 1995 of reversed polarity excursion in underlying Norkilili Member of Masek Beds with Emperor subchron. Lower limit: correlation of top of Bed IV with Matuyama Brunhes boundary 465
  14. 14. Table 1, Group 1 Continued Site Specimen Archaeology Date Method Selected references 466 OH 23, continued 990–370 ka Upper limit: 40Ar/39Ar Hay, 1976, 1990; Manega on Kerimasi caldera, 1993; Tamrat et al., 1995; probable source of Walter et al., 1991, 1992; Norkilili Member of Kimbel, 1995; White, 2000; Masek Beds. Lower Delson van Couvering, limit: correlation of 2000 normal polarity paleomagnetic zone at base of Masek Beds with Jaramillo subchron OH 28, WK, Olduvai Left innominate, femur Acheulian 780–620 ka (upper Bed K/Ar, paleomagnetism, Day, 1971, 1986; Leakey, Gorge, Tanzania IV) inferred sedimentation 1971a; Rightmire, 1979a; rates Leakey Hay, 1982; Hay, 1994 780 ka–1·2 Ma Paleomagnetism; This paper, based upon correlation of top of Tamrat et al., 1995 Bed IV with Matuyama Brunhes boundary 40 1·2–1·07 Ma Ar/39Ar, Hay, 1976, 1990; Manega, paleomagnetism; 1993; Tamrat et al., 1995; .   . .  correlation of normal Walter et al., 1991, 1992; polarity paleomagnetic Kimbel, 1995; White, 2000; zone at base of Masek Delson van Couvering, Beds with Jaramillo 2000 subchron OH 34, JK, Olduvai Femur and partial tibia Acheulian 0·8–1·1 Ma (Bed III) K/Ar, paleomagnetism Day, 1971; Leakey Roe, Gorge, Tanzania 1994; Hay, 1990, 1994 40 1·2–1·33 Ma Ar/39Ar, This paper, based upon paleomagnetism Hay, 1976, 1990; Tamrat et al., 1995; Walter et al., 1991, 1992; Kimbel, 1995; White, 2000
  15. 15. Table 1, Group 1 Continued Site Specimen Archaeology Date Method Selected references OH 51, GTC, Mandibular fragment None; Acheulian 0·8–1·1 Ma (Bed III) K/Ar, paleomagnetism Leakey Roe, 1994; Olduvai Gorge, elsewhere in Bed III Rightmire, 1990; Hay, Tanzania 1990, 1994 40 1·2–1·33 Ma Ar/39Ar, This paper, based upon paleomagnetism Hay, 1976, 1990; Tamrat et al., 1995; Walter et al., 1991, 1992; Kimbel, 1995; White, 2000 Saldanha (Hopefield Adult calvaria, Acheulian ca. 130–780 ka (Middle Associated fauna Drennan, 1953; Singer, Elandsfontein), South mandibular fragment Pleistocene) 1954; Cooke, 1963; Africa Partridge, 1982; Deacon, 1988 600–800 ka Associated fauna, cf Klein, 1973; Singer Bed IV Olduvai Gorge Wymer, 1968; Leakey Hay, 1982; Hay, 1994; Leakey Roe, 1994    ’ 500–200 ka Associated fauna Klein, 1988 700–400 ka Associated fauna Klein Cruz-Uribe, 1991; Klein, 1994; Rightmire, 1998 780 ka–1·2 Ma Associated fauna (cf This paper, based upon Olduvai Bed IV). Gentry, 1978; Klein Cruz Correlation of top of Uribe, 1991; White, 2000; Bed IV with Matuyama Tamrat et al., 1995; Walter Brunhes boundary et al., 1991, 1992; Kimbel, 1995 467
  16. 16. Table 1, Group 1 Continued 468 Site Specimen Archaeology Date Method Selected references Saldanha, continued 1·07–1·33 Associated fauna (cf This paper, based upon Olduvai Bed IV). Gentry, 1978; Klein Cruz Correlation of normal Uribe, 1991; White, 1999; polarity paleomagnetic Tamrat et al., 1995; Walter zone at base of Masek et al., 1991, 1992; Kimbel, Beds with Jaramillo 1995 subchron ´ Sale, Morocco Adult calvaria, cranial None 300 ka–1·0 Ma Associated fauna Jaeger, 1973, 1975; Howell, fragments, endocast 1978; Hublin, 1985, 1991, 1994; Dean et al., 1993 389–455 ka LU ESR on associated Hublin, 1991 bovid tooth enamel Sidi Abderrahman Partial mandible Acheulian Middle Pleistocene Geomorphology, Arambourg Biberson, (Casablanca) Morocco (‘‘Tensiftian’’=‘‘Riss’’) associated fauna 1956; Biberson, 1963; Howell, 1960, 1978 ´ Tighenif (formerly Three mandibles, Acheulian Middle Pleistocene Associated fauna Arambourg, 1955; Ternifine), Oran, parietal fragment Arambourg Hoffsteter, Algeria 1963; Howell, 1960, 1978; Balout et al., 1967; Tobias, .   . .  1968; Schwartz Tattersall, 2000 0·6–1·0 Ma Associated fauna Geraads, 1981; Jaeger, 1981 700 ka Associated fauna, Geraads et al., 1986 paleomagnetism Thomas 1 Quarry, Subadult partial None Middle Pleistocene Associated fauna, Ennouchi, 1969a, 1970; Morocco mandible, cranial and (‘‘Amirian’’) geomorphology Sausse, 1975b; Oakley et al., maxillary fragments 1977; Howell, 1960, 1978; Brauer, 1984a,b; Dean ¨ et al., 1993
  17. 17. Table 1, Group 1 Continued Site Specimen Archaeology Date Method Selected references ´ Wadi Dagadle, Maxilla, partial None 250 ka TL on basalt, de Bonis et al., 1984, 1988 Djibouti dentition associated fauna Group 2 (H. helmei or H. sapiens) Eliye Springs, West Adult cranium None None Surface find Brauer Leakey, 1986 ¨ Turkana, Kenya ES11693 14 Florisbad, South Adult cranium ?MSA? Infinite AAR, C Vogel Beaumont, 1972; Africa Bada et al., 1973 230 100 ka Th/234U, peat I, Dreyer, 1935, 1936; associated fauna Rightmire, 1978a; Clarke, 1985; Kuman Clarke, 1986; Brink, 1988; Butzer, 1988a; Kuman et al., 2000 260 ka ESR, direct assay on Grun et al., 1996 ¨ hominid tooth    ’ Guomde, Chari Fm, Adult cranium None 270–300 ka U-series, direct assays Brauer et al., 1992, 1997; ¨ Ileret, Kenya (KNM-ER 3884), on hominid cranium Feibel et al., 1989 femur (KNM-ER 999) femur 14 Haua Fteah, Libya Two young adult MSA (‘‘Levalloiso- Infinite C on burnt bone McBurney, 1961, 1967; mandibular fragments Mousterian’’) Tobias, 1967; Rak, 1998:364 130 Associated artefacts This paper, based upon (Generalized MSA is Debenath, 1994; Wendorf ´ stratified under Aterian et al., 1987, 1993a elsewhere in Sahara) 469
  18. 18. 470 Table 1, Group 2 Continued Site Specimen Archaeology Date Method Selected references Haua Fteah, Libya 90 ka Associated artefacts. This paper, based upon continued TL OSL reported Martini et al., 1996; for in situ Aterian Cremaschi et al., 1998 material in Libya; generalized MSA is stratified under Aterian elsewhere in Sahara 127–40 ka Associated artefacts Klein, 1999 14 Jebel Irhoud, Adult cranium (JI1), MSA (‘‘Levalloiso- Infinite C Ennouchi, 1966 Morocco adult calvaria (JI2), Mousterian’’) infant mandible (JI3), fragmentary postcranials (JI4) 90–125 ka (EU), 105 Extrapolation from Ennouchi, 1962, 1963, 190 ka (LU) ESR dates on 1968, 1969a,b; Howell, mammalian teeth 1978; Hublin, 1985, 1991, overlying in situ 1993; Hublin et al., 1987; hominid specimen JI4 Grun Stringer, 1991 ¨ .   . .  Mugharet el Aiya, Juvenile maxillary MSA, presumed Late Middle/Early Associated fauna Biberson, 1961, 1963; Morocco fragment with partial Aterian (not in situ) Late Pleistocene Debenath, 1980; Debenath ´ ´ dentition, adult tooth (‘‘Ouljian-Soltanian’’) et al., 1982, 1986; Amani Geraads, 1993; Hublin, 1993 60–90 ka Associated Aterian This paper, based upon artifacts; TL OSL Debenath, 1994; Martini ´ reported for in situ et al., 1996; Cremaschi Aterian material in et al., 1998 Libya
  19. 19. Table 1, Group 2 Continued Site Specimen Archaeology Date Method Selected references Ngaloba (Laetoli Adult cranium MSA 120 ka Correlation with Hay, 1976; Leakey Hay, Hominid 18) marker tuff in Lower 1982; Magori Day, 1983; Tanzania Ndutu Beds at Olduvai Day et al., 1980 Gorge bracketed by 14 C K/Ar dates 230 130–108 ka Th/234U on Hay, 1987 associated mammalian bone 100–200 ka Isoleucine Bada, 1987 epiminerization of associated mammalian bone 200 ka AAR on ostrich Manega, 1995 eggshell, correlation with units dated by 40 Ar/39Ar 40 200–370 ka Ar/39Ar, on Kerimasi This paper, based on Hay,    ’ caldera, probable 1976, 1990; Manega, 1993; source of Norkilili Tamra et al., 1995; Walter Member of Masek et al., 1991, 1992; Kimbel, Beds. 1995 40 200–490 ka Ar/39Ar, This paper, based on Hay, paleomagnetism; 1976, 1990; Manega, 1993; correlation of reversed Tamrat et al., 1995; Walter polarity excursion in et al., 1991, 1992; Kimbel, underlying Norkilili 1995; Delson van Member of Masek Couvering, 2000 Beds with Emperor subchron. 471
  20. 20. 472 Table 1, Group 2 Continued Site Specimen Archaeology Date Method Selected references 14 Omo II, Site PHS, Adult calvaria ?MSA? 39 ka C on Etheria shell in Day, 1969, 1972; Butzer, Kibish Formation, overyling unit 1969; Butzer et al., 1969; Ethiopia Merrick et al., 1973; Day Stringer, 1982 230 130 ka Th/234U on Etheria Day, 1969, 1972; Butzer, shell in overlying unit 1969; Butzer et al., 1969; Merrick et al., 1973; Day Stringer, 1982, 1991 Porc Epic (Dire u Mandibular fragment MSA Late Pleistocene Associated fauna Breuil et al., 1951; Vallois, Dawa), Ethiopia 1951; Clark, 1954, 1982; Brauer, 1984a; Howell, ¨ 1978 60–77 ka Obsidian hydration Clark et al., 1984; Clark, 1988 Singa, Sudan Calvaria ?MSA? Early Late Pleistocene Associated fauna Bate, 1951; Stringer, 1979; Brauer, 1984a,b; Stringer ¨ .   . .  et al., 1985; Clark, 1988 82–112 (EU), 133–187 ESR on associated Grun Stringer, 1991 ¨ (LU) mammalian teeth 190–130 ka U-series on calcrete McDermot et al., 1996 enclosing skull and associated mammalian teeth
  21. 21. Table 1 Continued Site Specimen Archaeology Date Method Selected references Group 3 (H. sapiens) Border Cave, South Adult calvaria (BC1), 2 MSA 90–115 ka Geomorphology, Cooke et al., 1945; Wells, Africa adult mandibles (BC 2 associated fauna, 1950, 1959; de Villiers, 5), infant partial extrapolation from 1973, 1978; Protsch, 1975; skeleton (BC3), adult overlying 14C dates, Beaumont et al., 1978; postcranial fragments amino acid racimzation Butzer et al., 1978; Rightmire, 1979b; Beaumont, 1980; Grun et al., ¨ 1990; Grun Stringer, ¨ 1991; Morris, 1992b; Miller et al., 1993, 1999; Pfeiffer Zehr, 1996; Pearson Grine, 1996 (but see Sillen Morris, 1996) 90–50 ka AAR, associated fauna, Klein, 1999 geomorphology, extrapolation from overlying 14C dates 14 Dar-es-Soltan, Cranial, maxillary MSA (Aterian) Infinite C McBurney, 1961;    ’ Morocco mandibular fragments Ferembach, 1976b; Howell, of 2 individuals, adult 1978; Brauer Rimbach, ¨ subadult 1990; Wendorf et al., 1990; Brauer, 1992; Debenath, ¨ ´ 1980, 1994; Debenath ´ et al., 1982, 1986; Hublin, 1993 60–90 ka Associated Aterian This paper, based upon artifacts; TL OSL Debenath, 1994; Martini ´ for in situ Aterian et al., 1996; Cremaschi material in Libya et al., 1998 473
  22. 22. 474 Table 1, Group 3 Continued Site Specimen Archaeology Date Method Selected references Die Kelders, South 24 teeth, mandibular MSA 40 ka, 60 ka (EU), ESR Grine Klein, 1985; Grine Africa fragment, 2 manual 80 ka (LU) et al., 1991; Avery et al., phalanges 1997 71–45 ka ESR, associated fauna, Klein, 1999 geologic context 60–80 ka ESR, TL, OSL, IRSL Grine, 2000; Schwarcz Rink, 2000; Feathers Bush, 2000 14 Equus Cave, South Mandibular left corpus MSA 93–44 ka U-series on tufa, C Grine Klein, 1985; Klein Africa (hyena layer) fragment with 2 on Mn patina et al., 1991 molars; additional isolated adult teeth 14 71–27 ka C, associated fauna, Klein, 1999 geologic context .   . .  Hoedjies Punt, South Cranial and postcranial MSA 70 ka U-series Volman, 1978; Berger Africa fragments, isolated Parkington, 1995 teeth 300–71 ka U-series, associated Klein, 1999 fauna, geologic context Kabua, Kenya Cranial mandibular Uncertain ?Late Pleistocene Geologic context Whitworth, 1966; fragments of 2 Rightmire, 1975 individuals

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