A multilevel analytical framework for studying cultural evolution in prehistoric hunter–gatherer societies

Over the past decade, a major debate has taken place on the underpinnings of cultural changes in human societies. A growing array of evidence in behavioural and evolutionary biology has revealed that social connectivity among populations and within them affects, and is affected by, culture. Yet the interplay between prehistoric hunter–gatherer social structure and cultural transmission has typically been overlooked. Interestingly, the archaeological record contains large data sets, allowing us to track cultural changes over thousands of years: they thus offer a unique opportunity to shed light on long‐term cultural transmission processes. In this review, we demonstrate how well‐developed methods for social structure analysis can increase our understanding of the selective pressures underlying cumulative culture. We propose a multilevel analytical framework that considers finer aspects of the complex social structure in which regional groups of prehistoric hunter–gatherers were embedded. We put forward predictions of cultural transmission based on local‐ and global‐level network metrics of small‐scale societies and their potential effects on cumulative culture. By bridging the gaps between network science, palaeodemography and cultural evolution, we draw attention to the use of the archaeological record to depict patterns of social interactions and transmission variability. We argue that this new framework will contribute to improving our understanding of social interaction patterns, as well as the contexts in which cultural changes occur. Ultimately, this may provide insights into the evolution of human behaviour.     

The evolution of altruistic social preferences in human groups

In this paper, we consider three hypotheses to account for the evolution of the extraordinary capacity for large-scale cooperation and altruistic social preferences within human societies. One hypothesis is that human cooperation is built on the same evolutionary foundations as cooperation in other animal societies, and that fundamental elements of the social preferences that shape our species'' cooperative behaviour are also shared with other closely related primates. Another hypothesis is that selective pressures favouring cooperative breeding have shaped the capacity for cooperation and the development of social preferences, and produced a common set of behavioural dispositions and social preferences in cooperatively breeding primates and humans. The third hypothesis is that humans have evolved derived capacities for collaboration, group-level cooperation and altruistic social preferences that are linked to our capacity for culture. We draw on naturalistic data to assess differences in the form, scope and scale of cooperation between humans and other primates, experimental data to evaluate the nature of social preferences across primate species, and comparative analyses to evaluate the evolutionary origins of cooperative breeding and related forms of behaviour.     

"To open oneself is a poor woman's trouble": embodied inequality and childbirth in South-Central Tanzania

Various theories exist for the ways in which social and material disparities are incorporated within human bodies and then expressed as health outcomes with uneven distributions. From a political economy perspective, one pathway involves processes of social exclusion that take place on articulating local and global fields of power. This study explores such situated processes as they produce and perpetuate embodied inequality at childbirth in the Kilombero Valley of South-Central Tanzania. Ethnographic narratives illustrate how these processes differentially affect the kind of care women seek and receive. Also described are women's complex yet pragmatic responses to potential exclusion in the attempt to secure a safe and otherwise positive outcome. In a culturally constructed world of childbirth, face-to-face claims on entitlement to biomedical services collide with enactments of discrimination at multiple levels, creating a space of contestation for social and material positioning as well as for physical well-being.     

Human Identity and the Evolution of Societies

Human societies are examined as distinct and coherent groups. This trait is most parsimoniously considered a deeply rooted part of our ancestry rather than a recent cultural invention. Our species is the only vertebrate with society memberships of significantly more than 200. We accomplish this by using society-specific labels to identify members, in what I call an anonymous society. I propose that the human brain has evolved to permit not only the close relationships described by the social brain hypothesis, but also, at little mental cost, the anonymous societies within which such alliances are built. The human compulsion to discover or invent labels to “mark” group memberships may originally have been expressed in hominins as vocally learned greetings only slightly different in function from chimpanzee pant hoots (now known to be society-specific). The weight of evidence suggests that at some point, conceivably early in the hominin line, the distinct groups composed of several bands that were typical of our ancestors came to be distinguished by their members on the basis of multiple labels that were socially acquired in this way, the earliest of which would leave no trace in the archaeological record. Often overlooked as research subjects, these sizable fission-fusion communities, in recent egalitarian hunter-gatherers sometimes 2,000 strong, should consistently be accorded the status of societies, in the same sense that this word is used to describe tribes, chiefdoms, and other cultures arising later in our history. The capacity of hunter-gatherer societies to grow sufficiently populous that not all members necessarily recognize one another would make the transition to larger agricultural societies straightforward. Humans differ from chimpanzees in that societal labels are essential to the maintenance of societies and the processes giving birth to new ones. I propose that anonymous societies of all kinds can expand only so far as their labels can remain sufficiently stable.     

How institutions shaped the last major evolutionary transition to large-scale human societies

What drove the transition from small-scale human societies centred on kinship and personal exchange, to large-scale societies comprising cooperation and division of labour among untold numbers of unrelated individuals? We propose that the unique human capacity to negotiate institutional rules that coordinate social actions was a key driver of this transition. By creating institutions, humans have been able to move from the default ‘Hobbesian’ rules of the ‘game of life’, determined by physical/environmental constraints, into self-created rules of social organization where cooperation can be individually advantageous even in large groups of unrelated individuals. Examples include rules of food sharing in hunter–gatherers, rules for the usage of irrigation systems in agriculturalists, property rights and systems for sharing reputation between mediaeval traders. Successful institutions create rules of interaction that are self-enforcing, providing direct benefits both to individuals that follow them, and to individuals that sanction rule breakers. Forming institutions requires shared intentionality, language and other cognitive abilities largely absent in other primates. We explain how cooperative breeding likely selected for these abilities early in the Homo lineage. This allowed anatomically modern humans to create institutions that transformed the self-reliance of our primate ancestors into the division of labour of large-scale human social organization.     

The evolution of the diversity of cultures

The abundant evidence that Homo sapiens evolved in Africa within the past 200,000 years, and dispersed across the world only within the past 100,000 years, provides us with a strong framework in which to consider the evolution of human diversity. While there is evidence that the human capacity for culture has a deeper history, going beyond the origin of the hominin clade, the tendency for humans to form cultures as part of being distinct communities and populations changed markedly with the evolution of H. sapiens. In this paper, we investigate 'cultures' as opposed to 'culture', and the question of how and why, compared to biological diversity, human communities and populations are so culturally diverse. We consider the way in which the diversity of human cultures has developed since 100,000 years ago, and how its rate was subject to environmental factors. We argue that the causes of this diversity lie in the distribution of resources and the way in which human communities reproduce over several generations, leading to fissioning of kin groups. We discuss the consequences of boundary formation through culture in their broader ecological and evolutionary contexts.     

Antiquity and Social Functions of Multilevel Social Organization Among Human Hunter-Gatherers

Three levels of social organization are recognized among human hunter-gatherers: the community, the domestic unit, and the band. We describe the key features of these three levels and show how they are intimately connected. We hypothesize that, in the course of human social evolution, bands emerged as a level of social organization within existing communities. As predators, hunter-gatherers live at lower population densities than chimpanzees (Pan troglodytes), even where the two species are sympatric. We propose that band formation evolved in humans from the more transient fissioning behavior seen within chimpanzee communities as a solution to the conflicting pressures of sustaining higher levels of cooperation required in hunting and the division of labor in a more dispersed community. If disputes break out, or if resources in the band territory are temporarily depleted, the existence of a wider community continues to be adaptive. To reconstruct the evolution of band society, we draw upon four lines of evidence: group (or network) size predicted from neocortex ratios, the distance materials were moved from their source during various periods in hominin evolution, ethnographic data on hunter-gatherer daily foraging ranges and population densities collated by the authors, and fossil hominin morphology. From these data, we conclude that key features of modern human hunter-gatherer social organization probably appear in the course of the evolution of Homo heidelbergensis.     

The ecology of social transitions in human evolution

We know that there are fundamental differences between humans and living apes, and also between living humans and their extinct relatives. It is also probably the case that the most significant and divergent of these differences relate to our social behaviour and its underlying cognition, as much as to fundamental differences in physiology, biochemistry or anatomy. In this paper, we first attempt to demarcate what are the principal differences between human and other societies in terms of social structure, organization and relationships, so that we can identify what derived features require explanation. We then consider the evidence of the archaeological and fossil record, to determine the most probable context in time and taxonomy, of these evolutionary trends. Finally, we attempt to link five major transitional points in hominin evolution to the selective context in which they occurred, and to use the principles of behavioural ecology to understand their ecological basis. Critical changes in human social organization relate to the development of a larger scale of fission and fusion; the development of a greater degree of nested substructures within the human community; and the development of intercommunity networks. The underlying model that we develop is that the evolution of ‘human society’ is underpinned by ecological factors, but these are influenced as much by technological and behavioural innovations as external environmental change.     

Natural pedagogy as evolutionary adaptation

We propose that the cognitive mechanisms that enable the transmission of cultural knowledge by communication between individuals constitute a system of 'natural pedagogy' in humans, and represent an evolutionary adaptation along the hominin lineage. We discuss three kinds of arguments that support this hypothesis. First, natural pedagogy is likely to be human-specific: while social learning and communication are both widespread in non-human animals, we know of no example of social learning by communication in any other species apart from humans. Second, natural pedagogy is universal: despite the huge variability in child-rearing practices, all human cultures rely on communication to transmit to novices a variety of different types of cultural knowledge, including information about artefact kinds, conventional behaviours, arbitrary referential symbols, cognitively opaque skills and know-how embedded in means-end actions. Third, the data available on early hominin technological culture are more compatible with the assumption that natural pedagogy was an independently selected adaptive cognitive system than considering it as a by-product of some other human-specific adaptation, such as language. By providing a qualitatively new type of social learning mechanism, natural pedagogy is not only the product but also one of the sources of the rich cultural heritage of our species.     

Landscapes of human evolution: models and methods of tectonic geomorphology and the reconstruction of hominin landscapes

This paper examines the relationship between complex and tectonically active landscapes and patterns of human evolution. We show how active tectonics can produce dynamic landscapes with geomorphological and topographic features that may be critical to long-term patterns of hominin land use, but which are not typically addressed in landscape reconstructions based on existing geological and paleoenvironmental principles. We describe methods of representing topography at a range of scales using measures of roughness based on digital elevation data, and combine the resulting maps with satellite imagery and ground observations to reconstruct features of the wider landscape as they existed at the time of hominin occupation and activity. We apply these methods to sites in South Africa, where relatively stable topography facilitates reconstruction. We demonstrate the presence of previously unrecognized tectonic effects and their implications for the interpretation of hominin habitats and land use. In parts of the East African Rift, reconstruction is more difficult because of dramatic changes since the time of hominin occupation, while fossils are often found in places where activity has now almost ceased. However, we show that original, dynamic landscape features can be assessed by analogy with parts of the Rift that are currently active and indicate how this approach can complement other sources of information to add new insights and pose new questions for future investigation of hominin land use and habitats.     

The scope of culture in chimpanzees, humans and ancestral apes

More studies have focused on aspects of chimpanzee behaviour and cognition relevant to the evolution of culture than on any other species except our own. Accordingly, analysis of the features shared by chimpanzees and humans is here used to infer the scope of cultural phenomena in our last common ancestor, at the same time clarifying the nature of the special characteristics that advanced further in the hominin line. To do this, culture is broken down into three major aspects: the large scale, population-level patterning of traditions; social learning mechanisms; and the behavioural and cognitive contents of culture. Each of these is further dissected into subcomponents. Shared features, as well as differences, are identified in as many as a dozen of these, offering a case study for the comparative analysis of culture across animal taxa and a deeper understanding of the roots of our own cultural capacities.     

Human social organization during the Late Pleistocene: Beyond the nomadic-egalitarian model

Many researchers assume that until 10–12,000 years ago, humans lived in small, mobile, relatively egalitarian bands. This “nomadic-egalitarian model” suffuses the social sciences. It informs evolutionary explanations of behavior and our understanding of how contemporary societies differ from those of our evolutionary past. Here, we synthesize research challenging this model and articulate an alternative, the diverse histories model, to replace it. We review the limitations of using recent foragers as models of Late Pleistocene societies and the considerable social variation among foragers commonly considered small-scale, mobile, and egalitarian. We review ethnographic and archaeological findings covering 34 world regions showing that non-agricultural peoples often live in groups that are more sedentary, unequal, large, politically stratified, and capable of large-scale cooperation and resource management than is normally assumed. These characteristics are not restricted to extant Holocene hunter-gatherers but, as suggested by archaeological findings from 27 Middle Stone Age sites, likely characterized societies throughout the Late Pleistocene (until c. 130 ka), if not earlier. These findings have implications for how we understand human psychological adaptations and the broad trajectory of human history.     

Universal and variable leadership dimensions across human societies

Many researchers have turned to evolutionary theory to better understand diversity in leadership. Evolutionary theories of leadership, in turn, draw on ethnographic cases of societies thought to more closely resemble the smaller-scale, face-to-face communities in which humans evolved. Currently, though, there is limited systematic data on the nature of leadership in such societies.We coded 109 dimensions of leadership, including costs and benefits relevant to evolutionary models, in 1212 ethnographic texts from 59 mostly nonindustrial populations in Human Relations Area Files (HRAF). We discovered evidence for both cultural universals in leadership, as well as important variation by continental region, subsistence strategy, group context, and leader sex. Candidate universals included that leaders were intelligent and knowledgeable, resolved conflicts, and received material and social benefits. Evidence for other leader dimensions varied by group context (e.g., there was more evidence that leaders of kin groups were older and tended to provide counsel and direction), subsistence (e.g., hunter-gatherers tended to lack leaders with coercive authority), and sex (e.g., female leaders tended to be associated with family contexts). There was generally more evidence of benefits than costs for both leaders and followers, with material, social, and mating benefits being particularly important for leaders, and material and other benefits important for followers.Shamans emerged as an important category of leaders who did not clearly conform to influential models that emphasize two leader strategies: using knowledge and expertise to provide benefits to followers vs. using physical formidability to impose costs. Instead, shamans and other leaders with supernatural abilities used their knowledge to both provide benefits and impose costs on others. We therefore propose a modified scheme in which leaders deploy their cognitive, social, material, and somatic capital to provide benefits and/or impose costs on others.     

Status competition,inequality, and fertility: implications for the demographic transition

The role that social status plays in small-scale societies suggests that status may be important for understanding the evolution of human fertility decisions, and for understanding how such decisions play out in modern contexts. This paper explores whether modelling competition for status—in the sense of relative rank within a society—can help shed light on fertility decline and the demographic transition. We develop a model of how levels of inequality and status competition affect optimal investment by parents in the embodied capital (health, strength, and skills) and social status of offspring, focusing on feedbacks between individual decisions and socio-ecological conditions. We find that conditions similar to those in demographic transition societies yield increased investment in both embodied capital and social status, generating substantial decreases in fertility, particularly under conditions of high inequality and intense status competition. We suggest that a complete explanation for both fertility variation in small-scale societies and modern fertility decline will take into account the effects of status competition and inequality.     

Innovation in the collective brain

Innovation is often assumed to be the work of a talented few, whose products are passed on to the masses. Here, we argue that innovations are instead an emergent property of our species'' cultural learning abilities, applied within our societies and social networks. Our societies and social networks act as collective brains. We outline how many human brains, which evolved primarily for the acquisition of culture, together beget a collective brain. Within these collective brains, the three main sources of innovation are serendipity, recombination and incremental improvement. We argue that rates of innovation are heavily influenced by (i) sociality, (ii) transmission fidelity, and (iii) cultural variance. We discuss some of the forces that affect these factors. These factors can also shape each other. For example, we provide preliminary evidence that transmission efficiency is affected by sociality—languages with more speakers are more efficient. We argue that collective brains can make each of their constituent cultural brains more innovative. This perspective sheds light on traits, such as IQ, that have been implicated in innovation. A collective brain perspective can help us understand otherwise puzzling findings in the IQ literature, including group differences, heritability differences and the dramatic increase in IQ test scores over time.     

Policing and punishment across the domains of social evolution

Several decades of research in humans, other vertebrates, and social insects have offered fascinating insights into the dynamics of punishment (and its subset, policing), but authors have only rarely addressed whether there are fundamental joint principles underlying the maintenance of these behaviors. Here we present a punisher/bystander approach rooted in inclusive fitness logic to predict which individuals should take on punishing roles in animal societies. We apply our scheme to societies of eusocial Hymenoptera and nonhuman vertebrate social breeders, and we outline potential extensions for understanding conflict regulation among cells in metazoan bodies and unrelated individuals in human societies. We highlight that: 1) no social unit is expected to express punishment behavior unless it collects positive inclusive fitness benefits that surpass alternative benefits of bystanding; 2) punishment with public good benefits can be maintained through either direct fitness benefits (coercion) or indirect fitness benefits (correction) or both; 3) differences across social systems in the distributions of power, relatedness, and reproductive options drive variation in the extent to which individuals actively punish; and 4) inclusive fitness logic captures many punishment‐relevant evolutionary and ecological variables in a single framework that appears to apply across very different types of social arrangements. Synthesis Researchers have long observed that individuals in animal societies punish (and by extension, police) each other, but they have rarely investigated whether general principles underlie this behavior across social arrangements. In this paper, we present a punisher/bystander approach rooted in inclusive fitness logic to predict which individuals should take on punisher roles in animal societies. We apply the approach to eusocial insects and cooperatively breeding vertebrates and outline extensions towards the control of cancer cell lineages and punishment in human groups. We highlight how variation in specific social variables may drive differences in punishing/policing across the social domains.     

Blood,War, and Ritual: Religious Ecology, ‘Strong’ Culture,and Human Sacrifice in the Premodern World

What social factors predict human sacrifice in premodern societies? After summarising key insights from competing theoretical perspectives that seek to explain the presence of human sacrifice in premodern societies, we empirically assess the explanatory utility of each theory. We draw from Stark's ‘moral communities’ argument and Alexander and Smith's insights regarding cultural autonomy to highlight how the macro-level organisation of premodern societies impacted the practice of human sacrifice. Using data from Murdock and White's Standard Cross-Cultural Sample, logistic regression models suggest that premodern societies that expressed community ties through religious ceremonies were more likely to engage in human sacrifice, while beliefs in spirit aggression are correlated with lower likelihood of human sacrifice. In terms of non-religious factors, societies that experienced frequent famine were slightly less likely to activity in the ancient world are partly a function of societal complexity. We conclude by specifying the theoretical implications of these findings.     

Kinship intensity and the use of mental states in moral judgment across societies

Decades of research conducted in Western, Educated, Industrialized, Rich, & Democratic (WEIRD) societies have led many scholars to conclude that the use of mental states in moral judgment is a human cognitive universal, perhaps an adaptive strategy for selecting optimal social partners from a large pool of candidates. However, recent work from a more diverse array of societies suggests there may be important variation in how much people rely on mental states, with people in some societies judging accidental harms just as harshly as intentional ones. To explain this variation, we develop and test a novel cultural evolutionary theory proposing that the intensity of kin-based institutions will favor less attention to mental states when judging moral violations. First, to better illuminate the historical distribution of the use of intentions in moral judgment, we code and analyze anthropological observations from the Human Area Relations Files. This analysis shows that notions of strict liability—wherein the role for mental states is reduced—were common across diverse societies around the globe. Then, by expanding an existing vignette-based experimental dataset containing observations from 321 people in a diverse sample of 10 societies, we show that the intensity of a society's kin-based institutions can explain a substantial portion of the population-level variation in people's reliance on intentions in three different kinds of moral judgments. Together, these lines of evidence suggest that people's use of mental states has coevolved culturally to fit their local kin-based institutions. We suggest that although reliance on mental states has likely been a feature of moral judgment in human communities over historical and evolutionary time, the relational fluidity and weak kin ties of today's WEIRD societies position these populations' psychology at the extreme end of the global and historical spectrum.     

Cladistic analyses of behavioural variation in wild Pan troglodytes: exploring the chimpanzee culture hypothesis

Long-term field studies have revealed considerable behavioural differences among groups of wild Pan troglodytes. Here, we report three sets of cladistic analyses that were designed to shed light on issues relating to this interpopulation variation that are of particular relevance to palaeoanthropology.In the first set of analyses, we focused on the proximate cause of the variation. Some researchers have argued that it is cultural, while others have suggested that it is the result of genetic differences. Because the eastern and western subspecies of P. troglodytes are well differentiated genetically while groups within the subspecies are not, we reasoned that if the genetic hypothesis is correct, the phylogenetic signal should be stronger when data from the eastern and western subspecies are analysed together compared to when data from only the eastern subspecies are analysed. Using randomisation procedures, we found that the phylogenetic signal was substantially stronger with in a single subspecies rather than with two. The results of the first sets of analyses, therefore, were inconsistent with the predictions of the genetic hypothesis.The other two sets of analyses built on the results of the first and assumed that the intergroup behavioural variation is cultural in nature. Recent work has shown that, contrary to what anthropologists and archaeologists have long believed, vertical intergroup transmission is often more important than horizontal intergroup transmission in human cultural evolution. In the second set of analyses, we sought to determine how important vertical transmission has been in the evolution of chimpanzee cultural diversity. The first analysis we carried out indicated that the intergroup similarities and differences in behaviour are consistent with the divergence of the western and eastern subspecies, which is what would be expected if vertical intergroup transmission has been the dominant process. In the second analysis, we found that the chimpanzee cultural data are not only comparable to a series of modern human cultural data sets in terms of how tree-like they are, but are also comparable to a series of genetic, anatomical, and behavioural data sets that can be assumed to have been produced by a branching process. Again, this is what would be expected if vertical inter-group transmission has been the dominant process in chimpanzee cultural evolution.Human culture has long been considered to be adaptive, but recent studies have suggested that this needs to be demonstrated rather than assumed. With this in mind, in the third set of analyses we investigated whether chimpanzee culture is adaptive. We found the hypothesis that chimpanzee culture is adaptive was supported by an analysis of data from the Eastern African subspecies, but not by an analysis of data from the eastern and western subspecies.The results of our analyses have implications for the number of subspecies in Pan troglodytes, the relationship between hominin taxa and Palaeolithic industries, and the evolution of hominin cognition and behaviour.     

Complex societies

The complexity of human societies of the past few thousand years rivals that of social insect societies. We hypothesize that two sets of social “instincts” underpin and constrain the evolution of complex societies. One set is ancient and shared with other social primate species, and one is derived and unique to our lineage. The latter evolved by the late Pleistocene, and led to the evolution of institutions of intermediate complexity in acephalous societies. The institutions of complex societies often conflict with our social instincts. The complex societies of the past few thousand years can function only because cultural evolution has created effective “work-arounds” to manage such instincts. We describe a series of work-arounds and use the data on the relative effectiveness of WWII armies to test the work-around hypothesis. Richerson received his Ph.D. degree in zoology from UC Davis in 1969. He is currently a professor in the Department of Environmental Science and Policy. In addition to his work in cultural evolution, he has worked on the limnology of Lake Tahoe and Clear Lake in California, and on Lake Titicaca in Peru and Bolivia. Boyd received his Ph.D. degree in ecology from UC Davis in 1975, though his thesis work was a resource economics problem. He is currently a professor of anthropology at UCLA. His research interests besides cultural evolution are game theory and a small bit of primatology from time to time.