Robert S. Walker

Co-Residence Patterns in Hunter-Gatherer Societies Show Unique Human Social Structure

Individuals in residential groups in contemporary hunter-gatherer societies are unrelated to each other. Contemporary humans exhibit spectacular biological success derived from cumulative culture and cooperation. The origins of these traits may be related to our ancestral group structure. Because humans lived as foragers for 95% of our species’ history, we analyzed co-residence patterns among 32 present-day foraging societies (total n = 5067 individuals, mean experienced band size = 28.2 adults). We found that hunter-gatherers display a unique social structure where (i) either sex may disperse or remain in their natal group, (ii) adult brothers and sisters often co-reside, and (iii) most individuals in residential groups are genetically unrelated. These patterns produce large interaction networks of unrelated adults and suggest that inclusive fitness cannot explain extensive cooperation in hunter-gatherer bands. However, large social networks may help to explain why humans evolved capacities for social learning that resulted in cumulative culture.

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.

Evolutionary History of Hunter-Gatherer Marriage Practices

Background The universality of marriage in human societies around the world suggests a deep evolutionary history of institutionalized pair-bonding that stems back at least to early modern humans. However, marriage practices vary considerably from culture to culture, ranging from strict prescriptions and arranged marriages in some societies to mostly unregulated courtship in others, presence to absence of brideservice and brideprice, and polyandrous to polygynous unions. The ancestral state of early human marriage is not well known given the lack of conclusive archaeological evidence. Methodology Comparative phylogenetic analyses using data from contemporary hunter-gatherers around the world may allow for the reconstruction of ancestral human cultural traits. We attempt to reconstruct ancestral marriage practices using hunter-gatherer phylogenies based on mitochondrial DNA sequences. Results Arranged marriages are inferred to go back at least to first modern human migrations out of Africa. Reconstructions are equivocal on whether or not earlier human marriages were arranged because several African hunter-gatherers have courtship marriages. Phylogenetic reconstructions suggest that marriages in early ancestral human societies probably had low levels of polygyny (low reproductive skew) and reciprocal exchanges between the families of marital partners (i.e., brideservice or brideprice). Discussion Phylogenetic results suggest a deep history of regulated exchange of mates and resources among lineages that enhanced the complexity of human meta-group social structure with coalitions and alliances spanning across multiple residential communities.

The trade-off between number and size of offspring in humans and other primates

Life-history theory posits a fundamental trade-off between number and size of offspring that structures the variability in parental investment across and within species. We investigate this ‘quantity–quality’ trade-off across primates and present evidence that a similar trade-off is also found across natural-fertility human societies. Restating the classic Smith–Fretwell model in terms of allometric scaling of resource supply and offspring investment predicts an inverse scaling relation between birth rate and offspring size and a −¼ power scaling between birth rate and body size. We show that these theoretically predicted relationships, in particular the inverse scaling between number and size of offspring, tend to hold across increasingly finer scales of analyses (i.e. from mammals to primates to apes to humans). The advantage of this approach is that the quantity–quality trade-off in humans is placed into a general framework of parental investment that follows directly from first principles of energetic allocation.

Nonlinear scaling of space use in human hunter–gatherers

Use of space by both humans and other mammals should reflect underlying physiological, ecological, and behavioral processes. In particular, the space used by an individual for its normal activities should reflect the interplay of three constraints: (i) metabolic resource demand, (ii) environmental resource supply, and (iii) social behaviors that determine the extent to which space is used exclusively or shared with other individuals. In wild mammals, there is an allometric scaling relation between the home range of an individual and its body size: Larger mammals require more space per individual, but this relation is additionally modified by productivity of the environment, trophic niche, sociality, and ability to defend a territory [Kelt DA, Van Vuren D (1999) Ecology 80: 337–340; Kelt DA, Van Vuren D (2001) Am Nat 157:637–645; Haskell JP, Ritchie ME, Olff H (2002) Nature 418:527–530; Damuth J (1987) Biol J Linn Soc 31:193–246; Damuth J (1981) Nature 290:699–700; and other previously published work]. In this paper we show how similar factors affect use of space by human hunter–gatherers, resulting in a nonlinear scaling relation between area used per individual and population size. The scaling exponent is less than one, so the area required by an average individual decreases with increasing population size, because social networks of material and information exchange introduce an economy of scale.

Lethal coalitionary aggression and long-term alliance formation among Yanomamö men

Significance Humans, like chimpanzees, engage in coalitionary violence: Members of both species coordinate lethal activity against conspecifics. The origin and adaptive functions of this behavior are poorly understood, and data from tribal populations are rare. We examine the composition of lethal coalitions from the Yanomamö, a tribal society in Amazonia. In contrast to chimpanzees, Yanomamö coalitions are composed of individuals from different lineages and natal communities. Many coalition partners are ideal marriage exchange partners. Men who kill together more often are more likely to live together in the same village later in life and to engage in marriage exchange. Our results highlight connections between coalitionary aggression and alliance formation and illuminate differences in social structure distinguishing humans from other primates. Some cross-cultural evidence suggests lethal coalitionary aggression in humans is the product of residence and descent rules that promote fraternal interest groups, i.e., power groups of coresident males bonded by kinship. As such, human lethal coalitions are hypothesized to be homologous to chimpanzee (Pan troglodytes) border patrols. However, humans demonstrate a unique metagroup social structure in which strategic alliances allow individuals to form coalitions transcending local community boundaries. We test predictions derived from the fraternal interest group and strategic alliance models using lethal coalition data from a lowland South American population, the Yanomamö. Yanomamö men who kill an enemy acquire a special status, termed unokai. We examine the social characteristics of co-unokais or men who jointly kill others. Analyses indicate co-unokais generally are (i) from the same population but from different villages and patrilines, (ii) close age mates, and (iii) maternal half-first cousins. Furthermore, the incident rate for co-unokai killings increases if men are similar in age, from the same population, and from different natal communities. Co-unokais who have killed more times in the past and who are more genetically related to each other have a higher probability of coresidence in adulthood. Last, a relationship exists between lethal coalition formation and marriage exchange. In this population, internal warfare unites multiple communities, and co-unokais strategically form new residential groups and marriage alliances. These results support the strategic alliance model of coalitionary aggression, demonstrate the complexities of human alliance formation, and illuminate key differences in social structure distinguishing humans from other primates.

Evolutionary history of partible paternity in lowland South America

Partible paternity, the conception belief that more than one man can contribute to the formation of a fetus, is common in lowland South America and characterized by nonexclusive mating relationships and various institutionalized forms of recognition and investment by multiple cofathers. Previous work has emphasized the fitness benefits for women where partible paternity beliefs facilitate paternal investment from multiple men and may reduce the risk of infanticide. In this comparative study of 128 lowland South American societies, the prevalence of partible paternity beliefs may be as much as two times as common as biologically correct beliefs in singular paternity. Partible paternity beliefs are nearly ubiquitous in four large language families—Carib, Pano, Tupi, and Macro-Je. Phylogenetic reconstruction suggests that partible paternity evolved deep in Amazonian prehistory at the root of a tentative Je-Carib-Tupi clade. Partible paternity often occurs with uxorilocal postmarital residence (males transfer), although there are exceptions. Partible paternity may have benefits for both sexes, especially in societies where essentially all offspring are said to have multiple fathers. Despite a decrease in paternity certainty, at least some men probably benefit (or mitigate costs) by increasing their number of extramarital partners, using sexual access to their wives to formalize male alliances, and/or sharing paternity with close kin.

Reports - Life-History Consequences of Density Dependence and the Evolution of Human Body Size

Previous attempts to explain variation in human growth and development emphasize the energetic constraints imposed by malnutrition and disease. However, this approach does not address the evolutionary effects of mortality risk on ontogenetic variation, a common theme in life‐history studies. The conventional approach can be reconciled with life‐history theory by considering the effect of mortality on the rates and timing of maturity in subsistence‐based human populations. Humans slow down growth and development and demonstrate smaller adult body sizes in high‐population‐density contexts, presumably because of increased nutritional constraints and disease loads. In addition, there is evidence of mortality‐based selection for relatively faster/earlier ontogeny in small‐bodied hunter‐gatherers living at high densities. This finding may be interpreted as an evolved reaction norm for earlier reproductive maturity and consequent smaller adult body size in high‐mortality regimes. In sum, comparative results support density‐dependent effects on body size that act through two pathways—nutritional constraints and juvenile mortality—at varying intensities, contributing to a nearly twofold range in body size across human societies.

Bayesian phylogeography of the Arawak expansion in lowland South America

Phylogenetic inference based on language is a vital tool for tracing the dynamics of human population expansions. The timescale of agriculture-based expansions around the world provides an informative amount of linguistic change ideal for reconstructing phylogeographies. Here we investigate the expansion of Arawak, one of the most widely dispersed language families in the Americas, scattered from the Antilles to Argentina. It has been suggested that Northwest Amazonia is the Arawak homeland based on the large number of diverse languages in the region. We generate language trees by coding cognates of basic vocabulary words for 60 Arawak languages and dialects to estimate the phylogenetic relationships among Arawak societies, while simultaneously implementing a relaxed random walk model to infer phylogeographic history. Estimates of the Arawak homeland exclude Northwest Amazonia and are bi-modal, with one potential homeland on the Atlantic seaboard and another more likely origin in Western Amazonia. Bayesian phylogeography better supports a Western Amazonian origin, and consequent dispersal to the Caribbean and across the lowlands. Importantly, the Arawak expansion carried with it not only language but also a number of cultural traits that contrast Arawak societies with other lowland cultures.

Population stability, cooperation, and the invasibility of the human species

The biogeographic expansion of modern humans out of Africa began ≈50,000 years ago. This expansion resulted in the colonization of most of the land area and habitats throughout the globe and in the replacement of preexisting hominid species. However, such rapid population growth and geographic spread is somewhat unexpected for a large primate with a slow, density-dependent life history. Here, we suggest a mechanism for these outcomes by modifying a simple density-dependent population model to allow varying levels of intraspecific competition for finite resources. Reducing intraspecific competition increases carrying capacities, growth rates, and stability, including persistence times and speed of recovery from perturbations. Our model suggests that the energetic benefits of cooperation in modern humans may have outweighed the slow rate of human population growth, effectively ensuring that once modern humans colonized a region long-term population persistence was near inevitable. Our model also provides insight into the interplay of structural complexity and stability in social species.