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- 1. Evolutionary Anthropology 109 ARTICLES On Stony Ground: Lithic Technology, Human Evolution, and the Emergence of Culture ´ ROBERT FOLEY AND MARTA MIRAZON LAHR Culture is the central concept of anthropology. Its centrality comes from the fact human evolution different and what it that all branches of the discipline use it, that it is in a way a shorthand for what is that it is necessary to explain. It is at makes humans unique, and therefore defines anthropology as a separate disci- once part of our biology and the thing pline. In recent years the major contributions to an evolutionary approach to that sets the limits on biological ap- culture have come either from primatologists mapping the range of behaviors, proaches and explanations. Just to among chimpanzees in particular, that can be referred to as cultural or “proto- add further confusion to the subject, it cultural”1,2 or from evolutionary theorists who have developed models to account is also that which is universally shared for the pattern and process of human cultural diversification and its impact on by all humans and, at the same time, human adaptation.3–5 the word used to demarcate differ- ences between human societies and groups. As if this were not enough for Theoretically and empirically, pa- that paleoanthropology can play in any hard-worked concept, it is both a leoanthropology has played a less the development of the science of cul- trait itself and also a process. When prominent role, but remains central to tural evolution. In particular, we want treated as a trait, culture can be con- the problem of the evolution of cul- to consider the way in which informa- sidered to be the trait or the means by ture. The gap between a species that tion from stone-tool technology can which that trait is acquired, transmit- includes Shakespeare and Darwin be used to map the pattern of cultural ted, changed, and used (that is, among its members and one in which evolution and thus throw light on the learned, taught, and socially passed a particular type of hand-clasp plays a nature of the apparent gap that lies on). It exists in the heads of humans major social role has to be significant. between humans and chimpanzees. and is manifested in the products of However, that gap is an arbitrary one, First, we discuss the various meanings actions. To add one further dimen- filled by the extinction of hominin of the culture concept and the role of sion, culture is seen by some as the species other than Homo sapiens. Pa- paleoanthropology in its use. Second, equivalent of the gene, and hence a leoanthropology has the potential to we look at how stone-tool technology particulate unit (the meme) that can fill that gap, and thus provide more of can be used to map cultural evolution be added together in endless permu- a continuum between humans and and provide insights into the cultural tations and combinations, while to other animals. Furthermore, it pro- capacities of different hominin spe- others it is as a large and indivisible vides the context, and hence the selec- cies. Third, we consider the inferences whole that it takes on its significance. tive environment, in which cultural that can be made from stone-tool In other words, culture is everything capabilities evolved, and so may pro- technology for the timing of major to anthropology, and it could be ar- vide insights into the costs and bene- events in cultural evolution. gued that in the process it has also fits involved in evolving cultural adap- become nothing.3,5–10 tations. The pervasive nature of the culture In this paper we focus on the role EVOLUTION, CULTURE AND concept means that evolutionary an- ANTHROPOLOGY thropology must also tackle the prob- lems it throws up. This is not the place Culture in Anthropology either to argue that the concept Robert Foley and Marta Mirazon Lahr are ´ Culture is the jam in the sandwich should be abandoned as of little or no at the Leverhulme Centre for Human Evo- of anthropology. It is all-pervasive. It analytical utility (one of us attempted lutionary Studies at the University of Cam- is used to distinguish humans from this several years ago, to no noticeable bridge, Downing Street, Cambridge. E-mail: raf10@cam.ac.uk and mbml1@cam.ac.uk apes (“everything that man does that effect11) nor to come up with a cut- the monkeys do not” (Lord Raglan)) ting-edge redefinition that will clear and to characterize evolutionarily de- away a century of obfuscation (we Evolutionary Anthropology 12:109 –122 (2003) rived behaviors in both living apes leave that in the capable hands of the DOI 10.1002/evan.10108 Published online in Wiley InterScience and humans. It is often both the ex- Editor of Evolutionary Anthropol- (www.interscience.wiley.com). planation of what it is that has made ogy). Rather, we wish to consider how
- 2. 110 Evolutionary Anthropology ARTICLES TABLE 1. Cultural Capacity Is Cognitively Based, but Is Correlated With a Number of Manifestations in the Realms of Learning, Social Organization, Symbolic Expression, and Patterns of Tradition. These Expressions May in Turn Be Visible in the Archeological and Fossil Record Broad Correlative Components of Culture Potential Paleobiological Manifestations Learning capacity Technology and technological variation Brain size? Social organization and structure Archeological density, structure and distribution Sexual dimorphism in fossil hominins Nonecologically functional elements of material culture Traits associated with symbolic thought Brain size? Anatomical basis for language Variation in material culture Tradition maintenance and change Regional variation and longevity of archeological components those aspects of anthropology that fore be a diachronic process. Compar- sistence through time). The possible deal with the deep past of the human isons between two living species, hu- paleobiological correlates of these are lineage—paleoanthropology—might mans and chimpanzees, can only shown in Table 1.12 throw light on the evolution of culture examine outcomes, not the actual pro- and the role it may have played in cess of transition. This must be in- EVOLUTIONARY HISTORY IN human evolution.12 ferred. The actual development of THE STONES: WHAT CAN THEY The problem in attempting this is more and more culture-bearing homi- that the sources of such evidence are nins must have occurred among spe- TELL US? limited, especially if little recourse is cies that are now extinct, to whom our We can now turn to stone technol- made either to analogical or phyloge- only access is through the fossil and ogy. From over two million years netic inferences from chimpanzees archeological record. lithic artifacts provide a rich and du- and other primates or extrapolation To search for something in the fos- rable source of information about the back from ethnography and psychol- sil and archeological records requires behavior of extinct hominins, and ogy. Paleoanthropology is limited to knowing what one is looking for, so thus greatly expand on the anatomical the archeological record for the evi- that a consideration of definitions of fossil evidence. The question to ask is dence it throws either on hominin culture cannot be entirely avoided. what sort of information can be de- cognition, and hence culture, or else Definitions of culture largely fall into rived from stone tools? on the cultural manifestations of be- two broad groups. Either they involve Archeologists have basically come havior. In practice, this means using the actual end products of behaviors up with two answers to this question. the record of stone tools, the primary that are inherently human (technol- On one hand, patterns in technology source of information about the be- ogy, for example) or they focus on the have been used to reconstruct popula- havior of prehuman hominins. processes that produce these out- tion histories, in a sense to construct comes—that is, the cognitive under- phylogenies of species and popula- pinnings. Most recent approaches tions (cultures, in other words). Stone Culture and have concentrated on the latter or, in tools were, in effect, treated as popu- Paleoanthropology other words, trying to get into the lation markers. This may be consid- There are two reasons why both minds of extinct species and popula- ered the phylogenetic and historical evolution and paleoanthropology are tions. This can only be done in terms approach. On the other hand, stone central to any discussion of culture. of correlates. Most definitions of cul- tools can be and have been interpreted The first is that the distinction be- ture involve three core elements: those as adaptive markers, often with little tween humans and other species is associated with learning, its depth or no phylogenetic signal, because usually drawn in some way around and extent, or the ability to acquire they are endlessly thrown up conver- the concept of culture—put simply, new information independent of a gently by the demands of the environ- we have it and they do not. Chimpan- tightly constrained genetic basis; ment and social organization, which zees chipping away at the margins of those associated with social organiza- thus reflect variability in behavioral tool making or grappling with the ru- tion and complexity; and those asso- response. This can be termed the diments of American sign language do ciated with symbolic thought, both its adaptive function approach. not really change this state of affairs. underlying cognitive basis and its By and large, these two approaches Given the fact that humans must have communication. In addition to these have been seen as alternatives, and to evolved from an acultural organism to core elements is the extent to which be in conflict with one another. Fur- one that possesses such capacities the behaviors derived from these ca- thermore, from a historical perspec- means that the evolution of culture is pacities are either capable of change tive, the adaptive function approach a major challenge to evolutionary the- and variability (a characteristic of cul- has generally supplanted and suc- ory. The second, related, aspect is that tural systems) and have a means of ceeded the phylogenetic and historical the evolution of culture must there- being maintained as traditions (per- approach, and has become the con-
- 3. ARTICLES Evolutionary Anthropology 111 sensus on which most Palaeolithic ar- stone tools were still seen as markers ings of the populations who made cheology operates. However, it is worth of peoples as they ebbed and flowed them, reflects the demands of the en- considering briefly the strengths and across the Palaeolithic landscape. The vironment and the responses of the weaknesses of each approach. high tide of the phylogenetic and his- populations to those demands within torical approach occurred when it was the constraints of raw-material avail- possible to draw simple boundaries ability. Phylogeny and History: around typological and technological The switch in emphasis was encap- Human Evolutionary History clusters and to associate them with sulated in the Mousterian debate of From Stone Tools cultural history and narrative. Thus, the 1970s, when Binford argued that the cultures of the Upper Palaeolithic, the variation in the frequencies of tool The idea that human evolutionary for example, were essentially analo- types in the rock shelters of the Dor- history might be reflected in stone- gous to ethnographic units, an anal- dogne and the Levant reflected differ- tool typology is one of the oldest in the ogy that was sometimes drawn all too ent activities being carried out, rather discipline and, in one form or an- explicitly.14 than the movements of different peo- other, has been a persistent theme This “from technology to culture to ple. The form of stone tools and their over the last one hundred and fifty people to history” approach has been frequencies in assemblages have been years or more. When Frere recognized subject to many criticisms, and is seen increasingly as the result of envi- the stone tools discovered in the eigh- largely associated with work by arche- ronmental and ecological demands teenth century as the product of hu- ologists in the first half of the twenti- and opportunities. Concomitant with mans, and at the same time recog- this view is the corollary that if the nized that they were very “primitive,” signal in the shapes of stone is func- he was drawing the first of many such conclusions. Stone-tool typology The idea of stone tools tion, it could not at the same time be phylogenetic and historical. could be seen to reflect the stages of as the markers of To this strongly ecological ap- human history, from the first simple chronology gradually proach has been added an additional flakes and cores through to the So- element, that of the constraints of lutrean points. During the first part of fell into disrepute, stone as a raw material and the pro- the twentieth century, this became especially as it was cess of knapping itself. It is clear that formalized in the schemes of Breuil, Burkitt, and Bordes.13 recognized that globally in some parts of the world good lithic materials are abundant, and in others The phylogenetic and historical ap- it was hard to maintain scarce. The strategies of stone tool proach generally encapsulated two basic components. The first was that if the model of universal manufacture would therefore be ex- pected to reflect this. The classic ex- stone tools were similar, then they stages and that there ample of this view has been the in- were made by the same sort of people, usually taken to mean people belong- was not necessarily any creasingly popular interpretation of the Movius Line as a raw material ing to the same culture, with greater chronological boundary within the Old World.17,18 or lesser implications for ethnic groups, depending on the time scale consistency to the The way in which stone tools are made—through a process of core and involved. The second was that the pattern of change. flake reduction—is also important. It level of sophistication or complexity has been argued that the differences of the tools reflected the cognitive or among typological elements are the cultural status of the population con- product of different degrees of reduc- cerned, usually more or less advanced tion, and that, for example, a few within the framework of the time. eth century. The move to a greater emphasis on adaptation and, more re- more blows and one type is trans- When these two components are put formed into another. Tool-type fre- together, one has a model for explain- cently, raw-material constraints, has greatly altered the way Palaeolithic ar- quency thus reflects use and the need ing prehistoric change in terms of the to retouch more or less. From an evo- movements of peoples through their cheology has been done and how the past is interpreted. lutionary perspective, the adaptive particular set of tools with a process function approach sees homoplasies of evolution toward greater cultural (convergent evolution brought about and, by implication, cognitive com- Adaptation and Function: through a combination of selection plexity. and constraints) as being rife, and The idea of stone tools as the mark- Information From Design therefore the phylogenetic signal of ers of chronology gradually fell into The alternative to the idea that stone tools as being very low. disrepute, especially as it was recog- stone tools reflect population and thus nized that globally it was hard to evolutionary history is that of adap- maintain the model of universal tive function, and is the consensus Back to Population History stages and that there was not neces- view of archeologists today.15,16 Vari- In recent years, however, there has sarily any chronological consistency ability in stone tools, rather than re- been a resurgence of interest in the to the pattern of change. Nonetheless, flecting the social and cultural group- interpretation of archeological mate-
- 4. 112 Evolutionary Anthropology ARTICLES rials in an evolutionary, in the sense of be expected to reside in the character- ideas we will develop here are one at- phylogenetic, perspective.19 This can istics of the various species and not to tempt at disentangling these signals. be seen in areas of direct interest to exhibit a great deal of sensitivity in re- First we look for the presence or ab- human evolution. One example is the lation to the environment. This allows sence of a correlation between biolog- association of the Aurignacian indus- us to consider whether stone-tool tech- ical evolution, based on morphologi- tries with the dispersals of modern nology covaries with phylogeny and cal affinities, and technological change, humans into Europe and, conversely, taxonomic status or with the environ- based on the distribution of technolog- the issue of whether there is a link ment, and thus provides an empirical ical modes. Second, we use this derived between Neanderthal populations and route into the problem. relationship to consider whether there the Mousterian in general and the In summary, therefore, we need to is an association between cultural out- Chatelperronean in particular.20,21 A consider stone tools both in an envi- put and the species involved, and where further example is the suggestion by ronmental context and in the context technological change occurs in relation Klein22 that the dispersal of Homo hei- of phylogeny. Both history and ecol- to biological change. Finally, we con- delbergensis into Europe is associated sider how these might relate to inferred with the Acheulean. We have also pro- cognition. Central to our argument is posed that stone tools are markers of that while environment is shaping the hominin geographical patterns,23 The extent to which technological demands, the nature of both in the long-term persistence of hominins’ behavioral response is cir- the Movius line and in the spread of hominins might have cumscribed by their cognitive abilities. Mode 3 or prepared core technologies possessed a greater or Thus, the link between technology and in Africa and Europe as part of a dis- persal of later archaic populations, as lesser degree of cultural phylogeny is crucial for determining the pattern of cultural evolution. well as modern humans.24 capacity might be The Evolution of Culture expected to be The Pattern of Hominin Evolution Through Stone Tools: Which reflected in the extent to To provide a framework, we can Approach? which we can see a briefly outline the pattern of hominin Given these two contrasting ap- good fit between the evolution from the origins of the ge- proaches to the information poten- tially locked in the stone tools, we ask environment and nus Homo. Figure 1 shows the distri- bution of proposed genus Homo taxa which one can give the most useful technology. Here the by time and geography. The earliest insights into the problem of the evo- proxy for culture is thus Homo, as well as the australo- lution of culture, and thus make use of pithecines, are excluded: Although the archeological record within the taken to be those there is clear evidence that they did field of anthropology more generally. aspects of the various make stone tools,25–27 this primarily Perhaps the common-sense answer is suggests either that Mode 1 technolo- the adaptive functional approach. definitions that gies are plesiomorphies of Homo, be- This would certainly be the preferred emphasize the ing developed among one or more option for most archeologists, as it australopithecine lineages, or else an represents the prevailing paradigm behavioral apomorphy at the base of Homo. The for the analysis of stone tool variation. manifestations of subsequent distribution of Mode 1 More importantly, as culture is pre- technologies shows the diversification sumably an adaptation, then it is only culture, variability, and and geographical radiation of the de- natural to use an adaptive approach to a high rate of change. scendants of Homo ergaster or possi- identify it in the past. The extent to bly earlier members of Homo. Among which hominins might have possessed these geographically widespread a greater or lesser degree of cultural members of Homo there appears to be capacity might be expected to be re- considerable diversity, with a distinc- flected in the extent to which we can ogy are important, as is the case in tive pattern to be found in Eastern see a good fit between the environ- most evolutionary problems. Testing Asia that has led some authorities to ment and technology. Here the proxy the various possibilities requires a du- distinguish between an African lin- for culture is thus taken to be those alistic approach. eage (H. ergaster) and an Asian one (H. aspects of the various definitions that erectus).28,29 emphasize the behavioral manifesta- STONE TOOL TECHNOLOGY Newer finds, such as those from tions of culture, variability, and a high Dmanisi,30 Ceprano,31,32 and Buia33 rate of change. AND HUMAN EVOLUTION support this perspective, although If, on the other hand, culture is seen Against this historical background, others such as the material from as a cognitive state reflecting the ability we propose that embedded in the Baka34 have been employed to ques- of the mind to generate new behaviors, Palaeolithic record are the signals of tion such a distinction. The evolution- then this can be something that might both adaptation and phylogeny. The ary changes that occur from a little
- 5. ARTICLES Evolutionary Anthropology 113 size different traits, including means of flake production, typological forms and frequencies, metrical variation, core reduction sequences, and mi- crowear patterns. The geographical and chronological scale of variation in each of these is very variable, and many show high levels of local, small- scale diversity rather than the large- scale one that we associate with homi- nin phylogeny biologically. We argue that in terms of mapping the general patterns of change and stability, what is needed is a scheme that operates on a global scale and reflects broad-scale change rather than local site varia- tion. To make a biological compari- son, we need a system that has high interpopulation variation relative to intrapopulation variation. Against this criterion, the most appropriate classification system is that of techno- logical modes, the major forms of lithic production (see Box 1). The principles have been developed elsewhere,23,24,39 but in brief consist in recognizing general technological traits and treating them cladistically. These traits refer to the basic means by which the stone tools were made and the broad nature of the artifactual outputs. Using Clark’s modes,40 five basic technologies have been recog- nized: Mode 1 being chopping tool and flake industries; Mode 2 being the production of bifaces and bifacially worked handaxes; Mode 3 being pre- Figure 1. Chronological and geographical distribution of recognized taxa of Homo. pared core technology; Mode 4 being lamellar or blade technology; and Mode 5 being microliths. Although there are more than 0.6 Myr have led to the present in Africa probably from continuities between them, they ex- view that there is a new taxon, H. hei- 150,000 years ago, but occur, presum- press more complex ways of making delbergensis, which had a larger cra- ably through population expansions, stone tools, leading toward greater con- nial vault and a generally more mod- in other parts of the world consider- trol and a more effective use of raw ern appearance, although retaining ably later: 100,000 years ago in West- material to produce particular end the extreme robusticity of the Lower ern Asia, 60,000 years ago in Austra- products. They are particularly suitable Pleistocene Homo species.35 This lia, and around 40,000 years ago in to be considered cladistically and so taxon is found in Africa and Europe, Mediterranean Europe and Eurasia.38 phylogenetically, because they are built and to some it may also be present in upon each other, and incorporate some East Asia. A further element of diver- Technological Modes, of the elements of “descent” that are sity can be added to this essentially Hominin Phylogeny, and the essential to an evolutionary approach. Middle Pleistocene pattern with H. It is important to emphasize that one Scale of Environmental antecessor, known from Spain.36 Fi- of the reasons that the technological nally, the terminal Middle Pleistocene Variation modes are appropriate for evolution- and the earlier parts of the Upper How do stone tools map on to a ary analysis is that they stress the Pleistocene show the evolution of two phylogeny of the genus Homo? This most derived elements in an assem- highly encephalized and derived raises the question of how we “mea- blage, for it is well-known that even forms of hominin, Neanderthals in sure” technological diversity. There is after the development of more derived Eurasia and modern humans in Af- no generally agreed means of doing modes, more “primitive” ones, in the rica.37 The latter, H. sapiens, are this, as different approaches empha- cladistic sense, persist.
- 6. 114 Evolutionary Anthropology ARTICLES Box 1. Clark’s Technological Modes Clark, working in the context of a to be relatively small compared to the variants. The Acheulean is known plethora of archeological cultures and size of the cores, and to lack, both on from Africa from dates close to 1.5 terminological diversity, attempted to the cores and the flakes, significantly million years ago, although it often is provide an overarching framework for invasive retouch. This mode resulted difficult to draw a line between this summarizing variation in Paleolithic in relatively little diversity of tool and the developed Oldowan Mode 1 and Mesolithic lithics on a global forms, relatively little by way of core industries. The bulk of well-docu- scale. He suggested that across the reduction, and lack of any prepara- mented African Acheulean sites are range of lithic assemblages there tion of the striking platforms. Mode 1 less than 1 million years old, and usu- could be seen some generalities that occurs extensively throughout the ally belong to the Middle Pleistocene. related to the way in which the stone Old World over much of the Pleisto- In Europe, Western Asia, and the In- tools were actually manufactured— cene and well into the Pliocene in dian subcontinent, the dated hence, the term modes. Clark had in sub-Saharan Africa. The African Acheulean sites fall mostly into the mind the technological modes of pro- Mode 1 industries are primarily Plio- Middle Pleistocene, although there duction for the Stone Age, upon cene and Lower Pleistocene, may be some evidence (at Ubeidiya) which were superimposed the variet- whereas in Eastern Asia they persist that it sporadically occurred earlier. ies brought about by cultural prefer- until the Upper Pleistocene. They also There has been prolonged contro- ence, economic need, and raw mate- occur in the Middle Pleistocene in Eu- versy over the presence of true Mode rial availability. His modes provide a rope. 2 industries in Eastern Asia. Although basic framework for grouping and Mode 2 saw the development of there is some evidence for bifacial separating stone-tool assemblages at two elements, although of course it stone tools in that region, there is a general rather than specific level. would have been possible for these to nothing truly like the recurrent They are described here in outline occur independently. The first of Acheulean of the west. It is this dis- form, with their broad geographical these was the ability to strike off rel- tinction that is represented by the and temporal distribution. atively large flakes so that they would Movius Line. Clark’s modes were based essen- have some of the size properties of Mode 3 represents a major shift in tially on the way in which the basic cores, but with a narrower cross-sec- the output of lithic production, al- flake-core relationship occurred. tional area, and thus be suitable for a though it shares with Mode 2 ele- Mode 1, comprising the Oldowan and greater amount of invasive retouch. It ments of the way tools are produced. Asian Pebble Tool and Chopping Tool was this that constituted the second The key difference is that the core is Traditions, constituted the simplest development, for it became possible prepared prior to striking off a major mode of production, the striking of a to retouch the resulting flakes in such flake as a means of having greater flake off a core. The number of flakes a way that secondary flakes were re- control over the shape and thickness could vary, but what held this system moved across the whole surface of of the flake. The actual means of of production together was the sim- the flake and on both sides. The result preparation, however, is probably ple platforms and lack of preparation was the bifacial tradition that is rep- similar to that used in the production involved. The flakes struck off tended resented by the Acheulean and its of handaxes. The outcome is a much
- 7. ARTICLES Evolutionary Anthropology 115 Box 1. Clark’s Technological Modes (continued) more diverse set of finished tools, elongated blades with narrow cross- flakes and blades that are retouched and hence a greater potential for vari- sections, which then are reworked and worked into various shapes in ability and a greater emphasis on extensively into diverse sets of sub- some contexts or are used as com- smaller items. Mode 3 constitutes the sidiary tool types. Although elongated posite unmodified tools in others. Mi- technologies of the European Middle flakes (that is, blades) are produced croliths are widely known in the later Palaeolithic and the African and In- by the Mode 3 technologies, the parts of prehistory. They form the ba- dian Middle Stone Age. Its presence Mode 4 system is different in that it is sis of the African Later Stone Age in the Middle Pleistocene of Eastern based on prismatic cores. Conven- from approximately 30,000 years Asia is disputed, but it may have had tionally, Mode 4 industries are asso- ago. However, there may have been a more extensive eastern distribution ciated with the Eurasian and North earlier occurrences of this mode (for in the Upper Pleistocene. African Upper Palaeolithic and occur example, the Howieson’s Poort in Mode 4 continues the trajectory of late (after 50,000 years) in the Upper southern Africa around 80,000 years Mode 3 in the sense that it is con- Pleistocene. Blades are also known ago). Microliths are also known in cerned with producing pieces off a to occur in earlier deposits, for exam- Southern Asia from around 30,000 core with the shape of those pieces ple in the Kapturin Beds in Kenya, years ago, more widely across Eu- being determined by the way in which and in the early Upper Pleistocene of rope and Asia in the latest parts of the the core has been prepared. In this Western Asia and Northern Africa, but Pleistocene, and in the early Holo- case, the preparation is designed to these are seldom prismatic. cene (the Mesolithic). Mode 5 indus- produce long flakes and results in cy- Mode 5 involves microlithic tech- tries are also known in the mid-Holo- lindrical prismatic cores and fine, nologies: the production of very small cene in Australia. Figure 2 shows the distribution of two trees is remarkably similar: Both rather than local scale of variation. the technological modes represented show deep African/Asian clades and These are, of course, two well-known in phylogenetic terms. It is perhaps relatively prolonged longevity of lin- and established facts, and it would striking that the overall shape of the eages. This confirms the continental perhaps be surprising if there was no
- 8. 116 Evolutionary Anthropology ARTICLES gence or at least a period of contact and cultural diffusion around 300 Ka. Elsewhere we have proposed that Ne- anderthals and modern humans may have shared a more recent Middle Pleistocene ancestor than H. heidel- bergensis, a population we named H. helmei.24 It should be noted that our use of H. helmei differs from that made later by McBrearty and Brooks48 to re- fer to the immediate African ancestor of modern humans only. The preceding evidence suggests that there is a strong but not entirely straightforward relationship between phylogeny and technological modes. This may seem to indicate that in terms of the two approaches dis- cussed earlier, the phylogenetic and historical approach is the most con- sistent with these data. This may sug- gest that there is not a strongly adap- tive element to technology. This is misleading in two ways. The first is that while the technology is adap- tive—that is, carrying out particular functions that enhance survivor- ship—it is strongly mediated by the cognitive capacities of those homi- nins, who appear to have been limited at least in terms of their ability to in- novate and vary their productions. This enhances the idea that the stone tools are providing insights into the evolution of the cognitive basis for culture. The second way in which we may be misled is if the approach Figure 2. Chronological and geographical distribution of lithic technologies in terms of through modes is insensitive to the modes. scale of variation that is significant at an adaptive level. This has been one of the criticisms leveled at the approach concordance given that they are sup- ones in the same region did—a mea- and can be discussed in terms of “pri- posedly the records of the same pop- sure of heritability, as it were, among vate histories.” ulations. How, though, does this pat- (admittedly nonreproducing) arti- tern relate to the expected scale of facts. There is a fidelity of form that Private Histories variation? The answer to this question defies the scale of ecological variation There are many caveats to the broad is that the scale observed seems to and seems to suggest that the varia- interpretation of the archeological reflect long-term phylogenetic pat- tion in stone tools as refuted in Clark’s record presented, of which the most terns more than fine-grained adaptive modes says more about the character- important one is that the modes ones. If environment was driving istics of their makers than the envi- clearly reflect only a small part of the Lower Paleolithic variability, one ronments in which they were living. variability in stone tools, and it could would perhaps expect a far more frag- There are, however, differences be- be argued that they are the only ones mented distribution, with, for exam- tween the two. Where most interpre- that reflect this scale of variation. Ty- ple, frequent oscillations between tations of the fossil phylogeny suggest pology, assemblage structure, and mi- Mode 1 and Mode 2 industries as hab- a divergence between European and crowear analysis might well display itats changed and as the availability of African lineages dating back to the diversity at either more general scales raw material varied from region to re- middle or early part of the Middle or more local ones. This in itself gion. This is not what is seen. Instead, Pleistocene, the shared technology of would not be surprising or necessarily the best predictor of what an artifact the Neanderthals and modern hu- a problem with this evolutionary his- is going to look like is what the earlier mans (Mode 3) suggests a later diver- tory model. Microwear,41 for example,
- 9. ARTICLES Evolutionary Anthropology 117 might well be expected to map onto a THE EVOLUTION OF CULTURE evidence for the origins of Homo. In very large scale of variability, as it is THROUGH HOMININ contrast, the earliest evidence for probably the case that different stone Homo ergaster does not relate to any EVOLUTION tools were used for the same purposes significant change in technology, but by different populations. In other Technology and Evolution: rather technological change occurs words, there is only one way to skin a Correlation and Causality considerably later, when Mode 2 ap- dead cat, but many tools that can be pears, after 1.4 Myr. It is also the case We can put this notion into practice used to do it. At the other extreme, the that Homo heidelbergensis, which is by considering the relationship be- detailed typological shape of the end- known from about 600,000 years ago, tween the major changes in modes product artifacts may well be ex- is not associated with a new techno- and the appearance of new taxa as pected to display local variation, as logical mode, although there is some shown in the fossil record.23,24,39 In these will be influenced both by the evidence to suggest that at this time Figure 3 a phylogeny for Homo is availability of raw materials and there is an intensification of biface shown, with the appearance and dis- small-scale cultural tradition, the Pa- production. Finally, when we look at leolithic equivalent of the different the later parts of human evolution, ways of hand-clasping or nest-build- there is some tentative reason for sug- ing found among chimpanzees. A pre- . . . although the modes gesting that the emergence of Mode 3 historic human example would be the differences in detailed bone harpoon do not tell the whole technologies in Africa may be associ- ated with a new morphology—what shape found among the epi-Paleo- story, they do tell an we have referred to elsewhere as lithic populations of northern Europe, important one. This Homo helmei. However, both H. sapi- which shared the same basic stone- ens and H. neanderthalensis make tool technology, and which Clark used might perhaps be a their appearance in the context of to identify social territories.42 pointer to the way in Mode 3 technologies, with Mode 4/5 We argue that although the modes only occurring tens of thousands of do not tell the whole story, they do tell which we think about years after the first anatomical evi- an important one. This might perhaps integrative approaches dence for modernity. be a pointer to the way in which we To many, the complexities of the think about integrative approaches to to human evolution. relationship between hominin lin- human evolution. There are many There are many sources eages and technology might lead to sources of information about the evo- of information about the the view that there is no relationship lutionary past, from fossils to archeol- at all. Certainly there is no simple ogy to genetics. While ultimately each evolutionary past, from causal relationship between the devel- must be the product of a single series fossils to archeology to opment of new technologies and spe- of historical events, nonetheless each ciation. There is not even a consistent may have to some extent a private his- genetics. While relationship, in the sense that techno- tory. Genes may record events that are ultimately each must be logical change always precedes ana- completely invisible archeologically— tomical change or vice versa. There is, indeed, one would expect them to— the product of a single nonetheless, an important pattern while the stone tools might be highly series of historical that requires explanation. What is sensitive to changes that are not seen in cranial morphology. Indeed, as the events, nonetheless likely is that different elements are re- lated to different events. Speciation number of genetic systems studied in- each may have to some or, more prosaically, the date of first creases, it is becoming clear that while extent a private history. appearances, is a demographic pro- they tell the same basic story, each cess, usually arising from the occur- one does have a private history: the Y rence of small isolated populations. It chromosome compared to mtDNA, is not inherent in this process that beta-globin compared to Alu inser- there should be a technological or be- tions, and so on. Different elements of appearance of the technological havioral or adaptive change. Rather, stone-tool technology may well also modes superimposed. It can be seen this process relates to genetic diver- have their own private histories, and that the relationship is far from gence, either through drift or selec- these histories may be regionally and straightforward (Fig. 4). It may be tion. The major behavioral changes chronologically specific. For this rea- that this complexity is at least partly that might be associated with any new son, technology may well not provide due to imprecise dating, but it may species could arise on either side of a single line of evidence and informa- also reflect to some extent the fact that that geographical boundary. Major tion, but separate ones relating to dif- while both the stones and the fossils adaptive changes, in other words, are ferent evolutionary events—some to tell the same story they are sensitive to not necessarily related to speciation. speciation, some to dispersals, some different parts of it. For example, we What they may be associated with are to behavioral grade shifts, some to can see that the emergence of stone- dispersals. That is, where technology cognition, some to ecology. tool technology predates the current confers a major adaptive advantage it
- 10. 118 Evolutionary Anthropology ARTICLES the Mode 2 industries, even subdi- vided into flake-based and nodule- based, are characteristic of particular periods and continents, and that they do not change much. It is perhaps a forgotten wonder of the archeological world that a French-trained archeolo- gist who knows of nothing but the Dordogne could go to the Cape of Good Hope and recognize the arti- facts and mode of production. What does this tell us about culture? Two things come to mind. The first is that across time there is clearly an increase in the complexity of the means by which tools are made, involving both more careful material selection, more forethought in the approach to pro- duction, and the potential for a greater diversity of outcomes. Unfash- ionable as it may be, this can be de- scribed as a progressive trend. How- ever, the question to ask is where across this trend are significant changes occurring. This is not just “chronological” variation. The modes persist much longer in some places than others (for example in Eastern Asia with Mode 1), suggesting that the evolution of the underlying cognitive capacities of the hominins was not uniform across the world. If the modes reflect culture or cultural ca- pacity, then culture is not evolving uniformly across the world’s hominin population. At present there is insuf- Figure 3. Comparison of chronological and geographical distribution of lithic modes and ficient data across Asia to understand Homo taxa. the details of this and whether it is a case of isolation or local selection, but as a problem it emphasizes the need leads to a geographical range expan- are deeply stable. Despite minor typo- to situate the archeological record on sion, and this will be visible: Hence logical variation and raw-material the hominin phylogeny. It is no longer the often apparently rapid widespread constraints, there is little doubt that possible to refer to generalized evolu- distributions of novel technologies.43 This may explain why the appearance of modern humans in Europe is asso- ciated with a new technology, the Au- rignacian or Upper Paleolithic, but the anatomical features associated with these populations have been present in Africa for as much as 150,000 years.44 The Evolution of Culture: Inferences From Technology What, though, can we learn about cultural evolution from modes? The Figure 4. Relationship between technological change and lineage change among hominins. most obvious point is that these tech- The left diagram shows the major lineages of Homo and where Mode changes occur within nological systems of fossil hominins them, while the right diagram shows how mode changes relate to “species” changes.
- 11. ARTICLES Evolutionary Anthropology 119 tion of cultural capacities within the cesses of demographic expansion into Between these two extremes lie genus Homo. various environments, and probably Modes 2 and 3. The development of The second cultural aspect is the reflect the processes described in Mode 2 at one level seems to show a stability of the modes across the Pleis- Shennan’s density model of cultural major change: the ability to strike off tocene, which has been extensively explosions.49 large flakes and invasively retouch discussed here and elsewhere. In one At the other end of the technological them in a controlled way, with a per- sense this stability mirrors a condition spectrum, the development of Mode 1 ception of the importance of final of culture—faithful replication of sys- technologies has been seen as a signif- shape.16,50 This shift occurs during tems— but it does so on a scale that is icant cultural evolutionary event, dis- the span of Homo ergaster. However, it manifestly very different from that of tinguishing more advanced hominins is worth noting several points about modern technologies. This argues ei- from apes.16 Although there is some the development of Mode 2. It does ther for a remarkable cultural tem- experimental evidence that chimpan- not appear with H. ergaster (1.8 Myr), plate beyond the capacities of modern zees are capable of stone fracture but several hundred thousand years humans or for the absence of another techniques, these appear to be later; while the end product (the cultural trait, the ability to innovate achieved with difficulty. Homo habi- Acheulean) is distinctive, it does and make modifications. This latter lis, or whichever Pliocene hominin merge more gradually with the Devel- possibility seems the more likely, with oped Oldowan (Mode 1); and there is a sense that one part of the cultural a considerable contrast between the program, imitation, was far more . . . there is some earlier forms and the later modern de- dominant in earlier hominins than it is in modern humans. As Byrne45 has experimental evidence rived Mode 2 that is associated with H. heidelbergensis, where there ap- shown for gorillas, imitation is quite a that chimpanzees are pears to be a much greater emphasis complex cognitive process, so this capable of stone on symmetry and regular form, espe- does not mean that these creatures cially once access was gained to the were not considerably more intelli- fracture techniques, flint sources of Europe. From the per- gent and culturally competent than these appear to be spective of cultural evolution, Mode 2 living apes. does represent a major cognitive shift, Finally, with regard to modes, we achieved with difficulty. but its full impact is a gradual pro- can ask whether the points at which Homo habilis, or cess, not a sudden punctuated event they change are significant events in followed by prolonged equilibrium. the evolution of culture or are what whichever Pliocene This seems to suggest that although has been referred to earlier as private hominin first made stone the rate of change is glacial in com- histories acting independently of the tools, was clearly able parison to modern cultural change, it rest of the hominin evolutionary does show a pattern of development record. The mode change that has at- to replicate the process that can be interpreted in terms of the tracted the most attention recently is consistently. This does refinement of a practice. that of Mode 4, the blade technology Mode 3 represents a different situa- associated with the Upper Palaeo- probably represent a tion. It can be cogently argued that the lithic.22,44,46 This has been strongly as- significant change in the basic technique of Mode 3, the prepa- sociated with the appearance of mod- ration of the core prior to flaking, is ern human behavior and the Out-of- process of cultural inherent in the Mode 2 technologies, Africa model of recent human evolution. and a “Levallois component” has long evolution. However, as various au- been recognized as a part of many thors have pointed out,24,47,48 it is dif- Acheulean assemblages. This has led ficult to pinpoint a direct cognitive some to suggest that the distinction change with this Mode. First, it is too between the two is insignificant. How- regionally specific, essentially being first made stone tools, was clearly able ever, although the actual technologi- confined to Eurasia. Second, it is too to replicate the process consistently. cal aspects of change may be contin- late, having occurred well after the This does probably represent a signif- uous, the outcomes are radically first appearance of modern humans icant change in the process of cultural different. Rather than the repetitive and after the diversification of the hu- evolution. Strout and coworkers have and monomorphic production of man population. If it was a cognitively used PET scans of people carrying out handaxes, instead there is the diver- and biologically based cultural shift, stone knapping to explore the cogni- sity of flake forms. The shift repre- then it occurred only after the major tive processes involved and shown sents a major change in the way stone populations of the world had sepa- that these do share similarities with cores (even if the cores are large rated, and therefore could not be a cognitive responses to tasks of a cul- flakes) are used and developed: They universal trait of humanity. Mode 4 tural nature in the extent to which become the template from which di- and, we argue, Mode 5 as well, are they coordinate motor control with versity can be produced rather than important, not as markers of major other aspects of cognition, especially the end product themselves. This can cognitive evolution, but of the pro- spatial processing. be seen in the increase in variation
- 12. 120 Evolutionary Anthropology that occurs in the Middle Stone Age Modes 4 and 5. Certainly, it seems and the earlier hominins. Further- and Middle Palaeolithic, both within that there is a strong contrast in be- more, the fact that these changes oc- and between assemblages.51,52 With havior and apparent cognitive flexibil- cur across the time of the lineages Mode 3 we see something that begins ity between hominins prior to H. hei- concerned suggests that this is not a to approach the variation we would delbergensis and those after. It is case of behavioral or cognitive sta- associate with modern cultural behav- perhaps significant that this is also the bility. The development of Mode 3 ior, and its appearance may be related period when brain-size evolution ac- (H. helmei, H. neanderthalensis, and to other substantive changes in behav- celerated. H. sapiens) represents an even greater ior.53 shift, with both standardization of Who Has Culture, Whatever form and diversification of end prod- Cultural Status of Extinct uct. Comparison across Modes 2 and That Is? 3 suggests that there is an earlier cog- Hominins nitive shift related to the ability to im- We are aware that compared to the On the basis of the preceding dis- rich tapestry of culture in the other itate and to maintain content and cussion, we could argue that the tech- papers in this issue, our version is form (tradition?), and a later one as- nological modes do provide useful in- somewhat stony and bare. There is no sociated with innovation. This latter sights into the evolution of culture, web of kinship or devious monkeys, change, when viewed in the context of but for this to be strengthened it needs no language, and no symbolic the evolution of modern humans and to be more firmly rooted into other thought. In a way, our intent has been the amazing accretion of diversity of aspects of human evolution. We have to trace the most basic of patterns in material culture that occurs through shown (Figs. 1–3) that there is consid- as broad a comparative context as the last 100,000 years, suggests that erable congruence in the broad distri- possible, so that we can see how cog- the evolution of these cultural capabil- bution of modes and hominin popula- nitive state might map on to the radi- ities was not a single event, but cumu- tions, but this is far from simple, and ation of hominins as seen in the fossil lative. Perhaps the most important that not all mode transitions show the record. This has meant confining our- conclusion is one that stresses the im- same pattern in relation to biological selves to a single source of informa- portance of looking at evolution diach- evolution. This is summarized in Ta- tion, stone tools, and a large-scale ap- ronically: The evolution of culture is not ble 2. proach, technological modes. Given a single step. Rather, the gap between From this two major points emerge. this limited approach, we can see that humans and chimpanzees, between a The first is that there is no simple re- the similarities between the fossil few termites for lunch and Beethoven, lationship between modes and homi- record and the technological one sug- is filled with incremental steps. nin species. For example, most tech- gest that the latter has a strong phylo- While it has been possible to gain nology-using hominins made Mode 1, genetic signal, and that this can be insights into the cognitive states of ex- itself an interesting insight into the interpreted as showing that the ability tinct hominins via the relationship be- evolution of culture, suggesting a deep to generate technological solutions to tween technological modes and mor- plesiomorphic conservatism for most adaptive problems was limited in phological affinities, it may be of human evolution. It is likely that many species. questioned how far we have demon- the origins of each mode lie in one If we return to the larger questions strated the absence or existence of lineage: Mode 1, an australopithecine?; relating to the evolution of culture culture. On one hand, it may be ar- Mode 2, H. ergaster; Mode 3, H. from the common ancestor with gued that as all the hominins make helmei; Modes 4 and 5, Homo sapiens. chimpanzees to modern humans, we and use stone tools, they are culture- It is clear, however, that these lineages can consider which among the many bearing; on the other hand, some all diversified into a number of de- species of hominins can be said to might say that as we have no access to scendent populations that persisted in have possessed culture or, more accu- symbolic thought or language, there is making the same stone tools. If these rately, how they compared in their no evidence for culture. In other are species, then speciation was not cultural capacities with either chim- words, the final interpretation de- the product of any technologically in- panzees or modern humans. The cul- pends on the definition of culture. The duced development. Indeed, in terms tural capacities of those hominins problem is how to proceed out of the of evolutionary process, it seems that making Mode 1 alone (early Homo, definitional problem. technologies change during the course early H. ergaster, and H. erectus) could One way is to recognize that culture of a lineage’s existence. be seen as very close to that of chim- is neither an absolute, present-or-ab- The second point is that if the stron- panzees in terms of their limited con- sent trait nor an indivisible whole. It is gest evidence for the evolution of en- trol and formalization of functional made up of a series of potentialities, hanced cultural capacities and their output, although the ability to gener- largely resting in cognition, and de- underlying cognition comes with free- ate standardized stone tools seems to pending on different mental thoughts. dom from the constraints of the envi- represent some sort of shift (perhaps We can differentiate, for example, be- ronment (and, in the case of technol- shared with some australopithecines?). tween imitation and copying as one ogy, this is presumably raw-material Those making Mode 2 (H. ergaster and element, which forms the basis for so- constraints), then this occurs in a se- H. heidelbergensis) show in the stan- cial transmission, social learning, and ries of stages during the development dardization of form and the remark- the maintenance of traditions, and in- of later Mode 2, more fully in Mode 3, able stability of tradition a consider- novation and elaboration, which and certainly with the elaboration of able difference from the chimpanzee forms the basis for cultural diversifi-
- 13. Evolutionary Anthropology 121 TABLE 2. Nature and Implications of Changes in Technological Modes Change through Associated Transition Nature of change Nature of output time Cultural inferences hominins Mode 0 Extension from stone- Relatively few different Little, although the Hominins probably not robustus? 2 tool use to stone- forms, with little Developed very dissimilar from garhi? Mode 1 tool modification, formal shape Oldowan can be apes, but control habilis or extension of Regional variation seen as a move and foresight rudolfensis nonstone tool probably just raw- toward Mode 2 involved in consistent ergaster modification to material related and greater fracturing, choosing erectus stone control, but the raw materials, and antecessor rate of change deploying tools (100,000s of years) shows a difference is very slow from the capabilities of living apes Mode 1 Ability to strike off Relatively few forms, Considerable The emphasis in Mode ergaster 2 large flake blanks but these show signs change through 2 technology is on heidelbergensis Mode 2 from cobbles or to of a preferred shape, time that may be greater planning use large nodules in and often exhibit related both to and goal-directed ways that allow symmetry technical behavior associated invasive retouch on Regional variation is competence and with demand for both sides probably largely the demand for particular shapes determined by raw particular Some evidence for materials (flakes preferred shapes cultural variation on versus nodules) (symmetry) a large geographical scale (cleavers in Africa and India) Mode 2 Transformation of the Diverse, predetermined Some directional Clear evidence for helmei 2 planning involved flakes, often very change from cultural variants neanderthalensis Mode 3 in Mode 2 toward thin, with potential early generalized regionally. sapiens the preparation of for modifying MSA to later, but Evidence for greater the core to allow extensively into most of the planning and greater control of different tools interassemblage awareness of flake production (especially points) variation is the indirect outputs Regional variation may development of be increasingly local styles (MTS, associated with Stillbay, cultural patterns Chatelperronean) rather than raw Final Mode 3 in materials Europe undergoes major change Mode 3 Continuation of the Blade blanks for use as Major change Evidence for ethnic sapiens 2 strategy of composite tools and through time, marking by Mode 4/5 emphasis on flake for secondary although not in technology and rather than core shaping any unidirectional other elements of production, and Major regional way toward material culture predetermination variation goes greater technical Localized cultural of shape. But with beyond raw material competence or traditions and emphasis on constraints, and refinement variants are narrow flakes probably reflects endemic (blades), very thin active strategies of flakes, and use and cultural miniaturization preference (Mode 5, microliths) cation. Each of these, and many oth- is that these contrasts are extremes on can track a series of different trajecto- ers, can be considered as a more finely a continuous scale and, furthermore, ries, each of which contributes to the graded scale. Whiten, in this volume, that they can vary independently. In final outcome. Thus, although we may has proposed that culture be tackled this sense, we return to the position never be able to speak in absolute through the search for contrasting that while culture as the end product terms about the cultural status of ex- features, which would include such of evolution may be a qualitatively dif- tinct hominins, we may be able to things as the existence or absence of ferent “whole,” its evolution is best scale them relative to humans and traditions. Perhaps what the paleoan- treated in a more reductionist and chimpanzees, and also gain insights thropological perspective adds to this piecemeal manner.11,12 In this way we into the process. For this to occur,
- 14. 122 Evolutionary Anthropology ARTICLES however, it is necessary for archeolo- cal advances in honor of F. Clark Howell. Engle- middle Pleistocene: the role of Homo heidelber- wood Cliffs NJ: Prentice Hall. p 569 –596. gensis. Evol Anthropol 6:218 –227. gists to work more closely with the 19 O’Brien MJ, Lyman RL, Saab Y, Saab E, Dar- 36 deCastro JMB, Arsuaga JL, Carbonell E, Ro- biological record that lies at the heart went J, Glover DS. 2002. Two issues in archaeo- sas A, Martinez I, Mosquera M. 1997. A hominid of human evolutionary history. logical phylogenetics: taxon construction and from the lower Pleistocene of Atapuerca, Spain: outgroup selection. J Theor Biol 215:133–150. possible ancestor to Neanderthals and modern 20 Mellars P. 1993. Archaeology and the popula- humans. Science 276:1392–1395. REFERENCES tion-dispersal hypothesis of modern human ori- 37 Stringer CB. 1995. The evolution and distri- gins in Europe. In: Aitken MJ, Stringer CB, Mel- bution of later Pleistocene human populations. lars PA, editors. The origins of modern humans In: VrbaES Denton GH, Partridge TC, Burckle 1 Whiten A, Goodall J, McGrew WC, Nishida T, and the impact of chronometric dating. Prince- LH, editors. Paleoclimate and evolution with em- Reynolds V, Sugiyama Y, et al. 1999. Cultures in ton: Princeton University Press. p 196 –216. phasis on human origins. New Haven: Yale Uni- chimpanzees. Nature 399:682–685. 21 Clark GA, Lindly JM. 1989. The case of con- versity Press. p 524 –531. 2 McGrew WC. 1992. Chimpanzee material cul- tinuity: observations on the biocultural transi- 38 Lahr MM, Foley RA. 1994. Multiple dispersals ture. Cambridge: Cambridge University Press. tion in Europe and Western Asia. In: Mellars P, and modern human origins. Evol Anthropol 3: 3 Cavalli-Sforza L, Feldman M. 1981. Cultural Stringer C, editors. The human revolution. Edin- 48 –60. transmission and evolution. Princeton: Princeton burgh: Edinburgh University Press. p 626 – 676. University Press. 39 Lahr MM, Foley RA. 2001. Mode 3, Homo 22 Klein RG. 1995. Anatomy, behaviour and Helmei, and the pattern of human evolution in 4 Boyd R, Richerson P. 1985. Culture and the modern human origins. J World Prehist 9:167– the Middle Pleistocene. In: Barham L, Robson evolutionary process. Chicago: University of Chi- 198. cago Press. Brown K, editors. Human roots: Africa and Asia 23 Foley RA. 1987. Hominid species and stone in the Middle Pleistocene. Bristol: Westbury 5 Laland KN, Odling-Smee J, Feldman MW. tools assemblages: how are they related? Antiq- Press. 2000. Niche construction, biological evolution, uity 61:380 –392. and cultural change. Behav Brain Sci 23:131– 40 Clark JGD. 1968. World prehistory: a new out- 24 Foley RA, Lahr MM. 1997. Mode 3 technolo- line, 2nd ed. Cambridge: Cambridge University 175. gies and the evolution of modern humans. Cam- Press. 6 Boesch C, Boesch H. 1983. Optimization of bridge Archaeol J 7:3–36. nut-cracking with natural hammers by wild 41 Keeley L, Toth N. 1981. Microwear polishes on 25 Leakey LSB. 1961. New finds at Olduvai early stone tools from Kenya. Nature 293:464 –465. chimpanzees. Behaviour 83:265–285. Gorge. Nature 189:649 –650. 7 Dawkins R. 1976. The selfish gene. London: 42 Clark JGD. 1972. The archaeology of stone 26 Susman RL. 1994. Fossil evidence for early age settlement. Ulster J Archaeol 35:3–16. Oxford University Press. hominid tool use. Science 265:1570 –1573. 8 Laland KN, Odling-Smee J, Feldman MW. 43 Lahr MM, Foley RA. 1998. Towards a theory 27 Asfaw B, White T, Lovejoy O, Latimer B, of modern human origins: geography, demogra- 2001. Cultural niche construction and human Simpson S, Suwa G. 1999. Australopithecus phy, and diversity in recent human evolution. evolution. J Evol Biol 14:22–33. garhi: a new species of early hominid from Ethi- Yearbook Phys Anthropol 41:137–176. 9 Lumsden CJ, Wilson EO. 1980. Translation of opia. Science 284:629 –635. epigenetic rules of individual behaviour into eth- 44 Klein RG. 1992. The archaeology of modern 28 Andrews PJ. 1984. An alternative interpreta- human origins. Evol Anthropol 1:5–14. nographic patterns. Proc Natl Acad Sci USA 77: tion of the characters used to define Homo erec- 4382–4386. 45 Byrne RW. 2002. Imitation of novel complex tus. Courier Forschunginstitut Seckenbergensis 10 Tomasello M. 2000. Culture and cognitive de- 69:167–175. actions: what does the evidence from animals velopment. Curr Dir Psychol Sci 9:37–40. mean? Adv Stud Behav 31:77–105. 29 Wood B. 1984. The origins of Homo erectus. 11 Foley RA. 1991. How useful is the culture Courier Foschungsinstitut Senkenberg 69:99 – 46 Klein RG. 2000. Archeology and the evolution concept in early human studies. In: Foley RA, 111. of human behavior. Evol Anthropol 9:17–36. editor. The origins of human behaviour. London: 47 Bar-Yosef O, Kuhn SL. 1999. The big deal 30 Lordkipanidze D, Vekua A. 2002. A new hom- Unwin Heinmann. p 25–38. about blades: laminar technologies and human inid mandible from Dmanisi (Georgia). J Hum 12 Foley RA. n.d. The emergence of culture in the Evol 42:A20 –A. evolution. Am Anthropol 101:322–338. context of hominin evolutionary patterns. In: 48 McBrearty S, Brooks AS. 2000. The revolution 31 Ascenzi A, Mallegni F, Manzi G, Segre AG, Levinson S, Jaisson P, editors. Evolution and that wasn’t: a new interpretation of the origin of Naldini ES. 2000. A re-appraisal of Ceprano cal- culture. Boston: MIT Press. In press. modern human behavior. J Hum Evol 39:453–563. varia affinities with Homo erectus, after the new 13 Bordes F. 1961. Typologie du paleolithique reconstruction. J Hum Evol 39:443–450. 49 Shennan SJ. 2000. Population, culture history ancien et modern. Bordeaux: Imprimeries Del- and the dynamics of culture change. Curr An- 32 Manzi G, Gracia A, Arsuaga JL. 2000. Cranial mas. thropol 41:811–835. discrete traits in the Middle Pleistocene humans 14 Bordes F. 1968. The old Stone Age. New York: from Sima de los Huesos (Sierra de Atapuerca, 50 Gamble C, Marshall G. 2001. A very remote McGraw Hill. Spain): does hypostosis represent any increase in period indeed. In: Milliken S, Cook J, editors. 15 Binford LR. 1973. Interassemblage variabili- “ontogenetic stress” along the Neanderthal lin- Papers on the Palaeolithic presented to Derek ty—the Mousterian and the functional argument. eage? J Hum Evol 38:425–446. Roe. Oxford: Oxbow Books. p 19 –27. In: Renfrew AC, editor. The exploration of cul- 33 Abbate E, Albianelli A, Azzaroli A, Benvenuti 51 Clark JD. 1992. African and Asian perspec- ture change. Pittsburgh: University of Pittsburgh M, Tesfamariam B, Bruni P, Cipriani N, Clarke tives on the origins of modern humans. Philos Press. p 227–254. RS, Ficcarelli G, Macchiarelli R, Napoleone G, Trans R Soc Series B 337:148 –178. 16 Schick K, Toth N. 1993. Making silent stones Papini M, Rook L, Sagri M, Tecle TM, Torre D, 52 McNabb J. 1997. The Neanderthal legacy: an speak. New York: Simon and Schuster. Villa I. 1998. A one-million-year-old Homo cra- archaeological perspective from Western Eu- 17 Pope GG. 1988. Recent advances in Far East- nium from the Danakil (Afar) Depression of Eri- rope—Mellars, P. J Archaeol Sci 24:575–576. ern paleoanthropology. Ann Rev Anthropol 17: trea. Nature 393:458 –460. 53 Stiner MC, Munro ND, Surovell TA, Tchernov 43–77. 34 Asfaw B, Gilbert WH, Beyene Y, Hart WK, E, Bar-Yosef O. 1999. Paleolithic population 18 Schick K. 1994. The Movius Line reconsid- Renne PR, WoldeGabriel G, Vrba ES, White TD. growth pulses evidenced by small animal exploi- ered: perspectives on the earlier Palaeolithic of 2002. Remains of Homo erectus from Bouri, Mid- tation. Science 283:190 –194. Asia. In: Corruccini RS, Ciochon RL, editors. In- dle Awash, Ethiopia. Nature 416:317–320. tegrative paths to the past: palaeoanthropologi- 35 Rightmire GP. 1998. Human evolution in the © 2003 Wiley-Liss, Inc.