Fission-fusion and the evolution of hominin social systems

The course of hominin evolution has involved successive migrations towards higher absolute latitudes over the past three million years. Poorer habitat quality further from the equator has led to the necessity for groups occupying higher latitudes to live at lower population densities. Coupled with a trend towards increasing group size over this time period, this tendency towards expansion has led to exponential increases in the area requirements of hominin groups, and a concomitant need to adjust foraging patterns. The current analyses suggest that the development of increasingly complex, multi-level fission-fusion social systems could have freed hominins of the foraging constraints imposed by large group sizes and low population densities. Analyses of the fossil record suggest latitudinally-driven differences in area requirements of the australopithecines from East and South Africa, and African and Asian Homo erectus. In contrast, chronologically-driven differences appear between H. erectus as a whole and Homo heidelbergensis, and between H. heidelbergensis and the Neanderthals. These results are discussed in relation to studies of the foraging patterns of primates and hunter-gatherers.     

Mutualism and manipulation in Hadza–honeyguide interactions

We investigated the ecology and evolution of interspecific cooperation between the Greater Honeyguide bird, Indicator indicator, and human hunter-gatherers, the Hadza of northern Tanzania. We found that honeyguides increased the Hadza's rate of finding bee nests by 560%, and that the birds led men to significantly higher yielding nests than those found without honeyguides. We estimate that 8–10% of the Hadza's total diet was acquired with the help of honeyguides. Contrary to most depictions of the human-honeyguide relationship, the Hadza did not actively repay honeyguides, but instead, hid, buried, and burned honeycomb, with the intent of keeping the bird hungry and thus more likely to guide again. Such manipulative behavior attests to the importance of social intelligence in hunter-gatherer foraging strategies. We present an evolutionary model for human-honeyguide interactions guided by the behavioral ecology of bees, non-human primates, and hunter-gatherers.     

The behavioral ecology of modern hunter-gatherers, and human evolution

Modern day hunter-gatherers are an obvious source of information about human life in the past. But can modern people really tell us anything about other hominids, those represented only in the fossil record? In a world of state governments and a global economy, can present-day foragers even tell us much about life before agriculture? Some behavioral ecologists think so. Their findings show (1) that foraging practices are closely related to the character and distribution of local resources, (2) that men, women and children react to foraging opportunities quite differently, and (3) that sex and age difference in these reactions have important social causes and consequences. Some results directly challenge long-held views about hunter-gatherer economics and social organization, and the scenarios of human evolution based on them.     

A Demographic Basis for Patrilineal Hordes

The evidence for patrilocal group organization among precontact hunter-gatherers is ambiguous. Observations among modern hunter-gatherers suggest that few, if any, are organized into patrilineal hordes or bands, but these observations stem from declining populations in transformed environments. These inadequacies in the data imbue the theoretical arguments concerning hunter-gatherer local organization with special importance. It is shown that ecological arguments against the viability of patrilineal bands fail because they conflate the membership of the camp with the band. The argument that variations in family size preclude patrilineal bands is found to be true in nonpolygynous populations. However, it is argued that patrilineal bands should develop in stable, strongly polygynous populations with small local groups, [hunter-gatherers, local organization, ecology, demography, polygyny]     

Society and territory in human evolution

A survey of recent data on the socio-territorial organization of primates, carnivores and human hunter-gatheres discloses some striking similarities among them. These common features are integrated into a theory of hominid social evolution. It is postulated that the hominids, throughout most of their evolution, were organized into stable groups with the capacity to disperse into largely independent subgroups that remain affiliated with each other. During the course of hominid evolution, territoriality became an increasingly important function of the larger, stable units. The analysis illustrates the value of combining data from primates, carnivores, and human hunter-gatherers in the reconstruction of early hominid behavior.     

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.     

Defining Higher Levels in the Multilevel Societies of Geladas (Theropithecus gelada)

Multilevel societies, identified by two or more nested levels (or modules) of organization, have been touted as some of the most complex social systems. However, few empirical studies have effectively quantified the association patterns that delineate the various levels in such societies. In particular, the multiple levels of gelada society were first described >3 decades ago, yet no operational definitions exist for the higher levels, i.e., levels above the one-male unit. In geladas, multiple units form aggregations that fission and fuse throughout the day, and throughout the year, blurring the distinctions between previously described higher social levels: teams, bands, and communities. Here we use 5?yr of data on the daily composition of a population of geladas living in the Simien Mountains National Park, Ethiopia to test the hypothesis that higher levels of gelada organization are discrete entities. If gelada aggregations are nothing more than a group of units that share a home range, then we expect a continuous distribution of unit association. If, however, gelada aggregations are indeed discrete organizational levels, then we expect discontinuity in the patterns of association. We found significant discontinuity at the 50% association level, indicating a sharp distinction between members of the same band (>50% association) and members of the same community (<50% association). We also found evidence that recently fissioned units form teams that associate significantly more than other band members. Thus, despite their extremely fluid social organization, gelada social levels are nevertheless clearly identifiable and quantifiable. Based on these results, we suggest that gelada society is an extremely flexible, multilevel society with fission–fusion dynamics, and as such gelada society presents an unusual example for understanding the evolution of modular societies.     

Variation in the Social Systems of Extant Hominoids: Comparative Insight into the Social Behavior of Early Hominins

The observed social systems of extant apes and humans suggest that the common ancestral state for Miocene hominoids was living in multimale–multifemale groups that exhibited a tendency to fission and fusion in response to ecological and/or social variables. The Hominoidea share a set of social commonalities, notably a social niche that extends beyond kin and beyond the immediate social group, as well as extensive intraspecific flexibility in social organization. We propose that an essential feature of hominoid evolution is the shift from limited plasticity in a generalized social ape to expanded behavioral plasticity as an adaptive niche. Whereas in most nonhominoid primates variability and flexibility take the shape of specific patterns of demographic flux and interindividual relationships, we can consider behavioral flexibility and plasticity as a means to an end in hominoid socioecological landscapes. In addition, the potential for innovation, spread, and inheritance of behavioral patterns and social traditions is much higher in the hominoids, especially the great apes, than in other anthropoid primates. We further suggest that this pattern forms a basis for the substantial expansion of social complexity and adaptive behavioral plasticity in the hominins, especially the genus Homo. Our objectives in this article are threefold: 1) summarize the variation in the social systems of extant hominoid taxa; 2) consider the evolutionary processes underlying these variations; and 3) expand upon the traditional socioecological model, especially with respect to reconstructions of early hominin social behavior. We emphasize a central role for both ecological and social niche construction, as well as behavioral plasticity, as basal hominoid characteristics. Over evolutionary time these characteristics influence the patterns of selection pressures and the resulting social structures. We propose that a mosaic of ecological and social inheritance patterns should be considered in the reconstruction of early hominin social systems.     

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.     

Social intelligence, human intelligence and niche construction

This paper is about the evolution of hominin intelligence. I agree with defenders of the social intelligence hypothesis in thinking that externalist models of hominin intelligence are not plausible: such models cannot explain the unique cognition and cooperation explosion in our lineage, for changes in the external environment (e.g. increasing environmental unpredictability) affect many lineages. Both the social intelligence hypothesis and the social intelligence-ecological complexity hybrid I outline here are niche construction models. Hominin evolution is hominin response to selective environments that earlier hominins have made. In contrast to social intelligence models, I argue that hominins have both created and responded to a unique foraging mode; a mode that is both social in itself and which has further effects on hominin social environments. In contrast to some social intelligence models, on this view, hominin encounters with their ecological environments continue to have profound selective effects. However, though the ecological environment selects, it does not select on its own. Accidents and their consequences, differential success and failure, result from the combination of the ecological environment an agent faces and the social features that enhance some opportunities and suppress others and that exacerbate some dangers and lessen others. Individuals do not face the ecological filters on their environment alone, but with others, and with the technology, information and misinformation that their social world provides.     

Autosomal Resequence Data Reveal Late Stone Age Signals of Population Expansion in Sub-Saharan African Foraging and Farming Populations

Background

A major unanswered question in the evolution of Homo sapiens is when anatomically modern human populations began to expand: was demographic growth associated with the invention of particular technologies or behavioral innovations by hunter-gatherers in the Late Pleistocene, or with the acquisition of farming in the Neolithic?

Methodology/Principal Findings

We investigate the timing of human population expansion by performing a multilocus analysis of≥20 unlinked autosomal noncoding regions, each consisting of ∼6 kilobases, resequenced in ∼184 individuals from 7 human populations. We test the hypothesis that the autosomal polymorphism data fit a simple two-phase growth model, and when the hypothesis is not rejected, we fit parameters of this model to our data using approximate Bayesian computation.

Conclusions/Significance

The data from the three surveyed non-African populations (French Basque, Chinese Han, and Melanesians) are inconsistent with the simple growth model, presumably because they reflect more complex demographic histories. In contrast, data from all four sub-Saharan African populations fit the two-phase growth model, and a range of onset times and growth rates is inferred for each population. Interestingly, both hunter-gatherers (San and Biaka) and food-producers (Mandenka and Yorubans) best fit models with population growth beginning in the Late Pleistocene. Moreover, our hunter-gatherer populations show a tendency towards slightly older and stronger growth (∼41 thousand years ago, ∼13-fold) than our food-producing populations (∼31 thousand years ago, ∼7-fold). These dates are concurrent with the appearance of the Late Stone Age in Africa, supporting the hypothesis that population growth played a significant role in the evolution of Late Pleistocene human cultures.     

Social learning among Congo Basin hunter-gatherers

This paper explores childhood social learning among Aka and Bofi hunter-gatherers in Central Africa. Existing literature suggests that hunter-gatherer social learning is primarily vertical (parent-to-child) and that teaching is rare. We use behavioural observations, open-ended and semi-structured interviews, and informal and anecdotal observations to examine the modes (e.g. vertical versus horizontal/oblique) and processes (e.g. teaching versus observation and imitation) of cultural transmission. Cultural and demographic contexts of social learning associated with the modes and processes of cultural transmission are described. Hunter-gatherer social learning occurred early, was relatively rapid, primarily vertical under age 5 and oblique and horizontal between the ages of 6 and 12. Pedagogy and other forms of teaching existed as early as 12 months of age, but were relatively infrequent by comparison to other processes of social learning such as observation and imitation.     

Hunter-Gatherers and the Origins of Religion

Recent studies of the evolution of religion have revealed the cognitive underpinnings of belief in supernatural agents, the role of ritual in promoting cooperation, and the contribution of morally punishing high gods to the growth and stabilization of human society. The universality of religion across human society points to a deep evolutionary past. However, specific traits of nascent religiosity, and the sequence in which they emerged, have remained unknown. Here we reconstruct the evolution of religious beliefs and behaviors in early modern humans using a global sample of hunter-gatherers and seven traits describing hunter-gatherer religiosity: animism, belief in an afterlife, shamanism, ancestor worship, high gods, and worship of ancestors or high gods who are active in human affairs. We reconstruct ancestral character states using a time-calibrated supertree based on published phylogenetic trees and linguistic classification and then test for correlated evolution between the characters and for the direction of cultural change. Results indicate that the oldest trait of religion, present in the most recent common ancestor of present-day hunter-gatherers, was animism, in agreement with long-standing beliefs about the fundamental role of this trait. Belief in an afterlife emerged, followed by shamanism and ancestor worship. Ancestor spirits or high gods who are active in human affairs were absent in early humans, suggesting a deep history for the egalitarian nature of hunter-gatherer societies. There is a significant positive relationship between most characters investigated, but the trait “high gods” stands apart, suggesting that belief in a single creator deity can emerge in a society regardless of other aspects of its religion.     

The complex structure of hunter-gatherer social networks

In nature, many different types of complex system form hierarchical, self-similar or fractal-like structures that have evolved to maximize internal efficiency. In this paper, we ask whether hunter-gatherer societies show similar structural properties. We use fractal network theory to analyse the statistical structure of 1189 social groups in 339 hunter-gatherer societies from a published compilation of ethnographies. We show that population structure is indeed self-similar or fractal-like with the number of individuals or groups belonging to each successively higher level of organization exhibiting a constant ratio close to 4. Further, despite the wide ecological, cultural and historical diversity of hunter-gatherer societies, this remarkable self-similarity holds both within and across cultures and continents. We show that the branching ratio is related to density-dependent reproduction in complex environments and hypothesize that the general pattern of hierarchical organization reflects the self-similar properties of the networks and the underlying cohesive and disruptive forces that govern the flow of material resources, genes and non-genetic information within and between social groups. Our results offer insight into the energetics of human sociality and suggest that human social networks self-organize in response to similar optimization principles found behind the formation of many complex systems in nature.     

从石制品技术特征视角探讨史前狩猎采集者的流动性

流动采食是狩猎采集者适应环境,获取生存资源的重要策略。对不同时空范围内狩猎采集群体的流动性具体特征进行探讨,有利于我们更加深入全面地理解史前狩猎采集者的人地互动模式、文化演进、社会转型等重要问题。本文以石制品技术组织理论为视角,结合相关研究案例,探讨如何通过分析考古出土的石制品特征,推断史前狩猎采集者不同形态特征的流动性策略,并讨论了通过石制品视角推断流动性特征的局限性及所需注意的事项。     

Parallel tool industries in New Caledonian crows

Individual specialization in the use of foraging tools occurs in hunter-gatherer societies but is absent in non-human primate tool use. 'Parallel tool industries' in hunter-gatherers are mainly based on strict sexual division of labour that is highly reliant on social conformity. Here, we show that 12 individuals in a population of New Caledonian crows on Maré Island had strong preferences for either stick tools or pandanus tools. Eight of the 12 crows had exclusive preferences. The individual specialization that we found is probably associated with different foraging niches. However, in spite of sexual size dimorphism there was no significant association between the sex of crows and their tool preferences. Our findings demonstrate that highly organized, strict sexual division of labour is not a necessary prerequisite for the evolution of parallel tool industries.     

A Papionin Multilevel Society as a Model for Hominin Social Evolution

Multilevel societies are unique in their ability to facilitate the maintenance of strong and consistent social bonds among some individuals while allowing separation among others, which may be especially important when social and sexual bonds carry significant and reliable benefits to individuals within social groups. Here we examine the importance of social and sexual bonds in the multilevel society of hamadryas baboons (Papio hamadryas) and apply these principles to social evolution in Plio-Pleistocene hominins. The behavior, adaptations, and socioecology of baboons (Papio spp.) have long been recognized as providing an important comparative sample to elucidate the processes of human evolution, and the social system of hamadryas baboons in particular shares even more similarities with humans than that of other baboons. Here we draw parallels between processes during the evolution of hamadryas social organization and those characterizing late Pliocene or early Pleistocene hominins, most likely Homo erectus. The higher costs of reproduction faced by female Homo erectus, exacerbated by an increased reliance on difficult to acquire, nutrient-dense foods, are commonly thought to have been alleviated by a strengthening of male–female bonds (via male provisioning and the evolution of monogamy) or by the assistance of older, postreproductive females (via grandmothering). We suggest that both of these social arrangements could have been present in Plio-Pleistocene hominins if we assume the development of a multilevel society such as that in hamadryas baboons. The evolution of a multilevel society thus underlies the adaptive potential for the complexity that we see in modern human social organization.     

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.     

Ardipithecus ramidus and the evolution of language and singing: An early origin for hominin vocal capability

In this paper we analyse the possibility that the early hominin Ardipithecus ramidus had vocal capabilities far exceeding those of any extant non-human primate. We argue that erect posture combined with changes in craniofacial morphology, such as reduced facial and jaw length, not only provide evidence for increased levels of pro-sociality, but also increased vocal ability. Reduced length of the face and jaw, combined with a flexed cranial base, suggests the larynx in this species was situated deeper in the neck than in chimpanzees, a trait which may have facilitated increased vocal ability. We also provide evidence that Ar. ramidus, by virtue of its erect posture, possessed a degree of cervical lordosis significantly greater than chimpanzees. This is indicative of increased mobility of the larynx within the neck and hence increased capacity to modulate vocalisations. In the paleoanthropological literature, these changes in early hominin skull morphology have to date been analysed in terms of a shift in mating and social behaviour, with little consideration given to vocally mediated sociality. Similarly, in the literature on language evolution there is a distinct lacuna regarding links between craniofacial correlates of social and mating systems and vocal ability. These are surprising oversights given that pro-sociality and vocal capability require identical alterations to the common ancestral skull and skeletal configuration. We therefore propose a model which integrates data on whole organism morphogenesis with evidence for a potential early emergence of hominin socio-vocal adaptations. Consequently, we suggest vocal capability may have evolved much earlier than has been traditionally proposed. Instead of emerging in the Homo genus, we suggest the palaeoecological context of late Miocene and early Pliocene forests and woodlands facilitated the evolution of hominin socio-vocal capability. We also propose that paedomorphic morphogenesis of the skull via the process of self-domestication enabled increased levels of pro-social behaviour, as well as increased capacity for socially synchronous vocalisation to evolve at the base of the hominin clade.