Neural correlates of Early Stone Age toolmaking: technology, language and cognition in human evolution

Archaeological and palaeontological evidence from the Early Stone Age (ESA) documents parallel trends of brain expansion and technological elaboration in human evolution over a period of more than 2Myr. However, the relationship between these defining trends remains controversial and poorly understood. Here, we present results from a positron emission tomography study of functional brain activation during experimental ESA (Oldowan and Acheulean) toolmaking by expert subjects. Together with a previous study of Oldowan toolmaking by novices, these results document increased demands for effective visuomotor coordination and hierarchical action organization in more advanced toolmaking. This includes an increased activation of ventral premotor and inferior parietal elements of the parietofrontal praxis circuits in both the hemispheres and of the right hemisphere homologue of Broca's area. The observed patterns of activation and of overlap with language circuits suggest that toolmaking and language share a basis in more general human capacities for complex, goal-directed action. The results are consistent with coevolutionary hypotheses linking the emergence of language, toolmaking, population-level functional lateralization and association cortex expansion in human evolution.     

Stone tools, language and the brain in human evolution

Long-standing speculations and more recent hypotheses propose a variety of possible evolutionary connections between language, gesture and tool use. These arguments have received important new support from neuroscientific research on praxis, observational action understanding and vocal language demonstrating substantial functional/anatomical overlap between these behaviours. However, valid reasons for scepticism remain as well as substantial differences in detail between alternative evolutionary hypotheses. Here, we review the current status of alternative 'gestural' and 'technological' hypotheses of language origins, drawing on current evidence of the neural bases of speech and tool use generally, and on recent studies of the neural correlates of Palaeolithic technology specifically.     

The Early Stone Age lithic assemblages of Gadeb (Ethiopia) and the Developed Oldowan/early Acheulean in East Africa

The Early Stone Age sites of Gadeb (Ethiopian South-East Plateau) were excavated under the direction of Desmond Clark in the 1970s. Dated to between 1.45 and 0.7 Ma, Gadeb proved that humans had already occupied high altitude areas in the Lower Pleistocene. Despite the importance of the Gadeb sites, their lithic assemblages were never published in detail, and no review of the stone tools has ever been reported since the original 1970s study. This paper updates the information available on Gadeb by presenting a systematic review of the lithic technology of several assemblages. The objectives are to evaluate the technological skills of Gadeb knappers and to contextualize them into the current discussion of the origins of the Acheulean and its possible coexistence with the so-called Developed Oldowan in East Africa.     

Behavioral inferences from Early Stone artifact assemblages: an experimental model

A methodological approach for assessing the nature and palaeographic distribution of early stone artifact assemblages is presented, modeled after an approach originally used for faunal analysis. By combining experimental replicative studies with careful analysis of Palaeolithic archaeological occurrences, it is potentially possible to reconstruct entire technological systems and to assess what stages of lithic reduction may be preferentially represented at high density artifact concentrations that we normally call archaeological “sites”, and what stages of reduction are found in lower density (“off site”) scatters.Based upon the results of this approach, alternative explanations for certain stages of lithic reduction being preferentially represented at a number of Plio-Pleistocene archaeological sites at Koobi Fora, Kenya are considered and evaluated with regard to early hominid organizational patterns. It appears that hominid stone technology was a relatively complex system by 1·9 to 1·4 million years B.P., involving significant transport and carrying of stone artifacts representing various stages of reduction and suggesting more foresight and planning than observed among extant nonhuman primates.The application of this approach to other Palaeolithic occurrences should enable anthropologists to obtain a better understanding of the organizational patterns of tool-making hominids throughout the course of human evolution.     

Modelling the evolution and diversity of cumulative culture

Previous work on mathematical models of cultural evolution has mainly focused on the diffusion of simple cultural elements. However, a characteristic feature of human cultural evolution is the seemingly limitless appearance of new and increasingly complex cultural elements. Here, we develop a general modelling framework to study such cumulative processes, in which we assume that the appearance and disappearance of cultural elements are stochastic events that depend on the current state of culture. Five scenarios are explored: evolution of independent cultural elements, stepwise modification of elements, differentiation or combination of elements and systems of cultural elements. As one application of our framework, we study the evolution of cultural diversity (in time as well as between groups).     

Generativity,hierarchical action and recursion in the technology of the Acheulean to Middle Palaeolithic transition: A perspective from Patpara,the Son Valley,India

The Acheulean to Middle Palaeolithic transition is one of the most important technological changes that occurs over the course of human evolution. Here we examine stone artefact assemblages from Patpara and two other excavated sites in the Middle Son Valley, India, which show a mosaic of attributes associated with Acheulean and Middle Palaeolithic industries. The bifaces from these sites are very refined and generally small, but also highly variable in size. A strong relationship between flake scar density and biface size indicates extensive differential resharpening. There are relatively low proportions of bifaces at these sites, with more emphasis on small flake tools struck from recurrent Levallois cores. The eventual demise of large bifaces may be attributed to the curation of small prepared cores from which sharper, or more task-specific flakes were struck. Levallois technology appears to have arisen out of adapting aspects of handaxe knapping, including shaping of surfaces, the utilization of two inter-dependent surfaces, and the striking of invasive thinning flakes. The generativity, hierarchical organization of action, and recursion evident in recurrent Levallois technology may be attributed to improvements in working memory.     

Modeling the formation of Early Stone Age artifact concentrations

The nature of stone artifact concentrations at early Plio-Pleistocene sites in East Africa is evaluated with regard to hominid transport behaviors responsible for their formation. These archaeological occurrences indicate ranging behaviors involving deliberate and repeated transport of flaked stone artifacts. The stone transported to archaeological sites within the time range of Homo habilis indicates planned transport of tools or material for tool manufacture to an extent far beyond transport behaviors reported among living apes, even stone hammer-using chimpanzees. Analysis of technological evidence in a lithic assemblage at a Plio-Pleistocene site at Koobi Fora (c. 1·5 ya) indicates on-site manufacturing activities and transport of flaked stone both to and from the site locale. Possible explanations for transport of stone artifacts are discussed in view of hominid strategies of environmental exploitation and resource utilization. A model is proposed for planned, habitual transport of artifacts by hominids positively correlated with distance of planned foraging range. In this model, larger-scale sites tended to develop at locales favorably located near abundant resources, where stone imports were high but export was relaxed due to the proximity of resources to be processed.     

Sleep intensity and the evolution of human cognition

Over the past four decades, scientists have made substantial progress in understanding the evolution of sleep patterns across the Tree of Life. 1 , 2 Remarkably, the specifics of sleep along the human lineage have been slow to emerge. This is surprising, given our unique mental and behavioral capacity and the importance of sleep for individual cognitive performance. 3 - 5 One view is that our species' sleep architecture is in accord with patterns documented in other mammals. 6 We promote an alternative view, that human sleep is highly derived relative to that of other primates. Based on new and existing evidence, we specifically propose that humans are more efficient in their sleep patterns than are other primates, and that human sleep is shorter, deeper, and exhibits a higher proportion of REM than expected. Thus, we propose the sleep intensity hypothesis: Early humans experienced selective pressure to fulfill sleep needs in the shortest time possible. Several factors likely served as selective pressures for more efficient sleep, including increased predation risk in terrestrial environments, threats from intergroup conflict, and benefits arising from increased social interaction. Less sleep would enable longer active periods in which to acquire and transmit new skills and knowledge, while deeper sleep may be critical for the consolidation of those skills, leading to enhanced cognitive abilities in early humans.     

Big brains, small worlds: material culture and the evolution of the mind

New developments in neuroimaging have demonstrated that the basic capacities underpinning human social skills are shared by our closest extant primate relatives. The challenge for archaeologists is to explain how complex human societies evolved from this shared pattern of face-to-face social interaction. We argue that a key process was the gradual incorporation of material culture into social networks over the course of hominin evolution. Here we use three long-term processes in hominin evolution-encephalization, the global human diaspora and sedentism/agriculture-to illustrate how the cultural transmission of material culture allowed the 'scaling up' of face-to-face social interactions to the global societies known today. We conclude that future research by neuroimagers and archaeologists will need to investigate the cognitive mechanisms behind human engagement with material culture as well as other persons.     

New thinking: the evolution of human cognition

Humans are animals that specialize in thinking and knowing, and our extraordinary cognitive abilities have transformed every aspect of our lives. In contrast to our chimpanzee cousins and Stone Age ancestors, we are complex political, economic, scientific and artistic creatures, living in a vast range of habitats, many of which are our own creation. Research on the evolution of human cognition asks what types of thinking make us such peculiar animals, and how they have been generated by evolutionary processes. New research in this field looks deeper into the evolutionary history of human cognition, and adopts a more multi-disciplinary approach than earlier 'Evolutionary Psychology'. It is informed by comparisons between humans and a range of primate and non-primate species, and integrates findings from anthropology, archaeology, economics, evolutionary biology, neuroscience, philosophy and psychology. Using these methods, recent research reveals profound commonalities, as well striking differences, between human and non-human minds, and suggests that the evolution of human cognition has been much more gradual and incremental than previously assumed. It accords crucial roles to cultural evolution, techno-social co-evolution and gene-culture co-evolution. These have produced domain-general developmental processes with extraordinary power-power that makes human cognition, and human lives, unique.     

Culturally transmitted paternity beliefs and the evolution of human mating behaviour

Recent anthropological findings document how certain lowland South American societies hold beliefs in 'partible paternity', which allow children to have more than one 'biological' father. This contrasts with Western beliefs in 'singular paternity', and biological reality, where children have just one father. Here, mathematical models are used to explore the coevolution of paternity beliefs and the genetic variation underlying human mating behaviour. A gene-culture coevolutionary model found that populations exposed to a range of selection regimes typically converge on one of two simultaneously stable equilibria; one where the population is monogamous and believes in singular paternity, and the other where the population is polygamous and believes in partible paternity. A second agent-based model, with alternative assumptions regarding the formation of mating consortships, broadly replicated this finding in populations with a strongly female-biased sex ratio, consistent with evidence for high adult male mortality in the region. This supports an evolutionary scenario in which ancestral South American populations with differing paternity beliefs were subject to divergent selection on genetically influenced mating behaviour, facilitated by a female-biased sex ratio, leading to the present-day associations of female control, partible paternity and polygamy in some societies, and male control, singular paternity and monogamy in others.     

The North African Middle Stone Age and its place in recent human evolution

The North African Middle Stone Age (NAMSA, ～300‐24 thousand years ago, or ka) features what may be the oldest fossils of our species as well as extremely early examples of technological regionalization and ‘symbolic’ material culture (d'Errico, Vanhaeren, Barton, Bouzouggar, Mienis, Richter, Hublin, McPherron, Louzouet, & Klein, 2009 ; Scerri, 2013a ; Richter, Grün, Joannes‐Boyau, Steele, Amani, Rué, Fernandes, Raynal, Geraads, Ben‐Ncer Hublin, McPherron, 2017 ). The geographic situation of North Africa and an increased understanding of the wet‐dry climatic pulses of the Sahara Desert also show that North Africa played a strategic role in continental‐scale evolutionary processes by modulating human dispersal and demographic structure (Drake, Blench, Armitage, Bristow, & White, 2011 ; Blome, Cohen, Tryon, Brooks, & Russell, 2012 ). However, current understanding of the NAMSA remains patchy and subject to a bewildering array of industrial nomenclatures that mask underlying variability. These issues are compounded by a geographic research bias skewed toward non‐desert regions. As a result, it has been difficult to test long‐established narratives of behavioral and evolutionary change in North Africa and to resolve debates on their wider significance. In order to evaluate existing data and identify future research directions, this paper provides a critical overview of the component elements of the NAMSA and shows that the timing of many key behaviors has close parallels with others in sub‐Saharan Africa and Southwest Asia.     

Review. Studying cumulative cultural evolution in the laboratory

Cumulative cultural evolution is the term given to a particular kind of social learning, which allows for the accumulation of modifications over time, involving a ratchet-like effect where successful modifications are maintained until they can be improved upon. There has been great interest in the topic of cumulative cultural evolution from researchers from a wide variety of disciplines, but until recently there were no experimental studies of this phenomenon. Here, we describe our motivations for developing experimental methods for studying cumulative cultural evolution and review the results we have obtained using these techniques. The results that we describe have provided insights into understanding the outcomes of cultural processes at the population level. Our experiments show that cumulative cultural evolution can result in adaptive complexity in behaviour and can also produce convergence in behaviour. These findings lend support to ideas that some behaviours commonly attributed to natural selection and innate tendencies could in fact be shaped by cultural processes.     

Culture and the evolution of human cooperation

The scale of human cooperation is an evolutionary puzzle. All of the available evidence suggests that the societies of our Pliocene ancestors were like those of other social primates, and this means that human psychology has changed in ways that support larger, more cooperative societies that characterize modern humans. In this paper, we argue that cultural adaptation is a key factor in these changes. Over the last million years or so, people evolved the ability to learn from each other, creating the possibility of cumulative, cultural evolution. Rapid cultural adaptation also leads to persistent differences between local social groups, and then competition between groups leads to the spread of behaviours that enhance their competitive ability. Then, in such culturally evolved cooperative social environments, natural selection within groups favoured genes that gave rise to new, more pro-social motives. Moral systems enforced by systems of sanctions and rewards increased the reproductive success of individuals who functioned well in such environments, and this in turn led to the evolution of other regarding motives like empathy and social emotions like shame.     

The Oldowan site Aïn al Fil (El Kowm,Syria) and the first humans of the Syrian Desert

At present, the oldest traces of human cultures are found in Eastern Africa. New discoveries set anew the questions about human and animal dispersal into Eurasia. For over 1.8 million years, humans have been present in the Levant. An extensive program of surveys and excavations in the Syrian Desert showed that this part of the world was a very ancient land of settlement. In Central Syria, the oldest site, Aïn al Fil in the region of El Kowm, was excavated in 2008 and 2010. The lithic industry in the lowest layer can be characterized by numerous unretouched flakes, pebble-tools and core-like artefacts. This assemblage is typical in a broad sense of archaic Palaeolithic the debitage of which corresponds to mode 1. From a techno-typological point of view, this industry tallies quite well with the so-called Oldowan stage. It shows remarkable similarities with the oldest African assemblages. From a chronologic point of view, these levels occur before three positive events in the Matuyama paleomagnetic sequence. It seems consistent to place the Oldowan sequence around 1.8 Ma BP within the Olduvai subchron or just before Olduvai/Matuyama reversal limit. Together with those of the neighbouring site Hummal, these levels would be the oldest traces of human presence ever found in Syria. In the Levant, the first humans not only occupied favourable zones but regularly ventured deep into less welcoming environments suggesting an astonishing flexibility in their behavioural and survival skills.     

Percussive technology in human evolution: an introduction to a comparative approach in fossil and living primates

Percussive technology is part of the behavioural suite of several fossil and living primates. Stone Age ancestors used lithic artefacts in pounding activities, which could have been most important in the earliest stages of stone working. This has relevant evolutionary implications, as other primates such as chimpanzees and some monkeys use stone hammer-and-anvil combinations to crack hard-shelled foodstuffs. Parallels between primate percussive technologies and early archaeological sites need to be further explored in order to assess the emergence of technological behaviour in our evolutionary line, and firmly establish bridges between Primatology and Archaeology. What are the anatomical, cognitive and ecological constraints of percussive technology? How common are percussive activities in the Stone Age and among living primates? What is their functional significance? How similar are archaeological percussive tools and those made by non-human primates? This issue of Phil. Trans. addresses some of these questions by presenting case studies with a wide chronological, geographical and disciplinary coverage. The studies presented here cover studies of Brazilian capuchins, captive chimpanzees and chimpanzees in the wild, research on the use of percussive technology among modern humans and recent hunter–gatherers in Australia, the Near East and Europe, and archaeological examples of this behaviour from a million years ago to the Holocene. In summary, the breadth and depth of research compiled here should make this issue of Philosophical Transactions of the Royal Society B, a landmark step forward towards a better understanding of percussive technology, a unique behaviour shared by some modern and fossil primates.     

The evolution of teaching

Teaching, alongside imitation, is widely thought to underlie the success of humanity by allowing high-fidelity transmission of information, skills, and technology between individuals, facilitating both cumulative knowledge gain and normative culture. Yet, it remains a mystery why teaching should be widespread in human societies but extremely rare in other animals. We explore the evolution of teaching using simple genetic models in which a single tutor transmits adaptive information to a related pupil at a cost. Teaching is expected to evolve where its costs are outweighed by the inclusive fitness benefits that result from the tutor's relatives being more likely to acquire the valuable information. We find that teaching is not favored where the pupil can easily acquire the information on its own, or through copying others, or for difficult to learn traits, where teachers typically do not possess the information to pass on to relatives. This leads to a narrow range of traits for which teaching would be efficacious, which helps to explain the rarity of teaching in nature, its unusual distribution, and its highly specific nature. Further models that allow for cumulative cultural knowledge gain suggest that teaching evolved in humans because cumulative culture renders otherwise difficult-to-acquire valuable information available to teach.     

The Oldowan industry of Peninj and its bearing on the reconstruction of the technological skills of Lower Pleistocene hominids

The Oldowan technology has traditionally been assumed to reflect technical simplicity and limited planning by Plio-Pleistocene hominids. The analysis of the Oldowan technology from a set of 1.6-1.4 Ma sites (ST Site Complex) in Peninj adds new information regarding the curated behavior of early hominids. The present work introduces new data to the few published monographic works on East African Oldowan technology. Its relevance lies in its conclusions, since the Peninj Oldowan assemblages show complex technological skills for Lower Pleistocene hominids, which are more complex than has been previously inferred for the Oldowan stone tool industry. Reduced variability of tool types and complex use of cores for flaking are some of the most remarkable features that identify the Oldowan assemblages from Peninj. Hominids during this period seem to have already been experimenting with pre-determination of the flaked products from cores, a feature presently assumed to appear later in time. Planning and template structuring of flaked products are integral parts of the Oldowan at Peninj.