Self-Interest and the Design of Rules

Rules regulating social behavior raise challenging questions about cultural evolution in part because they frequently confer group-level benefits. Current multilevel selection theories contend that between-group processes interact with within-group processes to produce norms and institutions, but within-group processes have remained underspecified, leading to a recent emphasis on cultural group selection as the primary driver of cultural design. Here we present the self-interested enforcement (SIE) hypothesis, which proposes that the design of rules importantly reflects the relative enforcement capacities of competing parties. We show that, in addition to explaining patterns in cultural change and stability, SIE can account for the emergence of much group-functional culture. We outline how this process can stifle or accelerate cultural group selection, depending on various social conditions. Self-interested enforcement has important bearings on the emergence, stability, and change of rules.     

Reservation food sharing among the Ache of Paraguay

We describe food transfer patterns among Ache Indians living on a permanent reservation. The social atmosphere at the reservation is characterized by a larger group size, a more predictable diet, and more privacy than the Ache typically experience in the forest while on temporary foraging treks. Although sharing patterns vary by resource type and package size, much of the food available at the reservation is given to members of just a few other families. We find significant positive correlations between amounts transferred among pairs of families, a measure of the "contingency" component required of reciprocal altruism models. These preferred sharing partners are usually close kin. We explore implications of these results in light of predictions from current sharing models. This research was supported by an L.S.B. Leakey Foundation grant and an NSF Graduate Fellowship to M. Gurven, and NSF Grant #9617692 to K. Hill and A. M. Hurtado. Michael Gurven recently obtained his Ph.D. from the University of New Mexico and is now an assistant professor at UC-Santa Barbara. His current interests include exploring ways in which socioecology influences variation in cooperation within and across human groups, and how cultural norms of fairness co-evolve with systems of resource production and distribution. Wesley Allen-Arave is pursuing his Ph.D. in anthropology at the University of New Mexico. His primary research interests focus on exploring variations across time and space in nonreciprocated altruistic acts, cooperation within social networks, and concerns over social approval. Kim Hill is a professor of anthropology in the Human Evolutionary Ecology (HEE) program at the University of New Mexico. His primary research interests include hunter-gatherer behavioral ecology, life history theory, food acquisition strategies, food sharing, cooperation, and biodiversity conservation in lowland South America. He has done fieldwork with Nahautl, Ache, Guarani, Hiwi, Mashco Piro, Matsiguenga, and Yora indigenous peoples of Central and South America. A. Magdalena Hurtado is associate professor of anthropology at the University of New Mexico. Her research interests include the evolution of cooperation between the sexes, infectious disease and immune system adaptations, the epidemiology of hunter-gatherer societies in transition, and the effects of health on economic productivity. During the past 20 years she has conducted fieldwork among several South American native populations but now works primarily among the Ache of eastern Paraguay.     

Our better nature: Does resource stress predict beyond-household sharing?

Food sharing and (to a lesser extent) labor sharing play central roles in the evolution of cooperation literature. One popular explanation for sharing beyond the family is that it reduces the likelihood of shortages by pooling risk across households. However, the frequency and scope of sharing have never been systematically documented across nonindustrial societies, and the literature is driven by theoretical models, experimental games, and case studies among a few extensively-studied populations. Here we explore the cross-cultural context, frequency, and scope of food and labor sharing customs in relation to resource stress. Using ethnographic data from a worldwide sample of 98 societies in the Standard Cross-Cultural Sample (SCCS), we test the following hypotheses: 1) customary sharing of food and labor beyond the household are cultural universals, 2) societies subject to more resource stress (including unpredictable food-destroying natural hazards) will share more frequently, and 3) the more frequent the resource stress, the broader the geographic and social scope of sharing customs. Hypotheses 1 and 2 are generally supported and are consistent with the theory that extensive beyond-household sharing is adaptive in societies that are subject to more resource stress. Hypothesis 3 was not supported and, contrary to our predictions, there is suggestive evidence that sharing beyond-relatives may be attenuated when resource stress is high. In light of these findings, we consider how resource stress may constitute an important selection pressure for maintaining extensive cooperation and help to explain the ubiquity of beyond-household sharing.     

Cultural evolution of cooperation: The interplay between forms of social learning and group selection

The role of cultural group selection in the evolution of human cooperation is hotly debated. It has been argued that group selection is more effective in cultural evolution than in genetic evolution, because some forms of cultural transmission (conformism and/or the tendency to follow a leader) reduce intra-group variation while creating stable cultural variation between groups. This view is supported by some models, while other models lead to contrasting and sometimes opposite conclusions. A consensus view has not yet been achieved, partly because the modelling studies differ in their assumptions on the dynamics of cultural transmission and the mode of group selection. To clarify matters, we created an individual-based model allowing for a systematic comparison of how different social learning rules governing cultural transmission affect the evolution of cooperation in a group-structured population. We consider two modes of group selection (selection by group replacement or by group contagion) and systematically vary the frequency and impact of group-level processes. From our simulations we conclude that the outcome of cultural evolution strongly reflects the interplay of social learning rules and the mode of group selection. For example, conformism hampers or even prevents the evolution of cooperation if group selection acts via contagion; it may facilitate the evolution of cooperation if group selection acts via replacement. In contrast, leader-imitation promotes the evolution of cooperation under a broader range of conditions.     

The evolution of daily food sharing: A Bayesian phylogenetic analysis

Some human subsistence economies are characterized by extensive daily food sharing networks, which may buffer the risk of shortfalls and facilitate cooperative production and divisions of labor among households. Comparative studies of human food sharing can assess the generalizability of this theory across time, space, and diverse lifeways. Here we test several predictions about daily sharing norms–which presumably reflect realized cooperative behavior–in a globally representative sample of nonindustrial societies (the Standard Cross-Cultural Sample), while controlling for multiple sources of autocorrelation among societies using Bayesian multilevel models. Consistent with a risk-buffering function, we find that sharing is less likely in societies with alternative means of smoothing production and consumption such as animal husbandry, food storage, and external trade. Further, food sharing was tightly linked to labor sharing, indicating gains to cooperative production and perhaps divisions of labor. We found a small phylogenetic signal for food sharing (captured by a supertree of human populations based on genetic and linguistic data) that was mediated by food storage and social stratification. Food sharing norms reliably emerge as part of cooperative economies across time and space but are culled by innovations that facilitate self-reliant production.     

The resilience of social norms of cooperation under resource scarcity and inequality — An agent-based model on sharing water over two harvesting seasons

Water governance remains a challenge for human societies, especially when the variation in resource inflow is large and the resource users are heterogeneous. We analyze with a coupled social-ecological systems (SES) model how socioeconomic and environmental changes affect the resilience of social norms governing resource use. In our model, agents have access to water as a common-pool resource and allocate it between rainy and dry seasons. While it is socially optimal to save water for the dry season, it is individually optimal to take water immediately. In our model, punishment of norm violators is the mechanism that may sustain cooperation. We show that the resilience of social norms could be affected by changes in socioeconomic and environmental conditions. Particularly, we find that social norms may collapse in times of resource scarcity and variability, especially if several drivers act in concert. Finally, we find that user heterogeneity in the form of different skills and inequality in land endowments may undermine cooperation. This implies that climatic changes and increased inequality – both potential drivers in the field – may affect community resilience and may lead to an erosion of social norms.     

Interethnic Interaction,Strategic Bargaining Power,and the Dynamics of Cultural Norms

Ethnic groups are universal and unique to human societies. Such groups sometimes have norms of behavior that are adaptively linked to their social and ecological circumstances, and ethnic boundaries may function to protect that variation from erosion by interethnic interaction. However, such interaction is often frequent and voluntary, suggesting that individuals may be able to strategically reduce its costs, allowing adaptive cultural variation to persist in spite of interaction with out-groups with different norms. We examine five mechanisms influencing the dynamics of ethnically distinct cultural norms, each focused on strategic individual-level choices in interethnic interaction: bargaining, interaction-frequency-biased norm adoption, assortment on norms, success-biased interethnic social learning, and childhood socialization. We use Bayesian item response models to analyze patterns of norm variation and interethnic interaction in an ethnically structured Amazonian population. We show that, among indigenous Matsigenka, interethnic education with colonial Mestizos is more strongly associated with Mestizo-typical norms than even extensive interethnic experience in commerce and wage labor is. Using ethnographic observations, we show that all five of the proposed mechanisms of norm adoption may contribute to this effect. However, of these mechanisms, we argue that changes in relative bargaining power are particularly important for ethnic minorities wishing to preserve distinctive norms while engaging in interethnic interaction in domains such as education. If this mechanism proves applicable in a range of other ethnographic contexts, it would constitute one cogent explanation for when and why ethnically structured cultural variation can either persist or erode given frequent, and often mutually beneficial, interethnic interaction.     

A Peak-Clustering Method for MEG Group Analysis to Minimise Artefacts Due to Smoothness

Magnetoencephalography (MEG), a non-invasive technique for characterizing brain electrical activity, is gaining popularity as a tool for assessing group-level differences between experimental conditions. One method for assessing task-condition effects involves beamforming, where a weighted sum of field measurements is used to tune activity on a voxel-by-voxel basis. However, this method has been shown to produce inhomogeneous smoothness differences as a function of signal-to-noise across a volumetric image, which can then produce false positives at the group level. Here we describe a novel method for group-level analysis with MEG beamformer images that utilizes the peak locations within each participant’s volumetric image to assess group-level effects. We compared our peak-clustering algorithm with SnPM using simulated data. We found that our method was immune to artefactual group effects that can arise as a result of inhomogeneous smoothness differences across a volumetric image. We also used our peak-clustering algorithm on experimental data and found that regions were identified that corresponded with task-related regions identified in the literature. These findings suggest that our technique is a robust method for group-level analysis with MEG beamformer images.     

The co-evolution of individual behaviors and social institutions

We present agent-based simulations of a model of a deme-structured population in which group differences in social institutions are culturally transmitted and individual behaviors are genetically transmitted. We use a standard extended fitness accounting framework to identify the parameter space for which this co-evolutionary process generates high levels of group-beneficial behaviors. We show that intergroup conflicts may explain the evolutionary success of both: (a) altruistic forms of human sociality towards unrelated members of one's group; and (b) group-level institutional structures such as food sharing which have emerged and diffused repeatedly in a wide variety of ecologies during the course of human history. Group-beneficial behaviors may evolve if (a) they inflict sufficient fitness costs on outgroup individuals and (b) group-level institutions limit the individual fitness costs of these behaviors and thereby attenuate within-group selection against these behaviors. Thus, the evolutionary success of individually costly but group-beneficial behaviors in the relevant environments during the first 90,000 years of anatomically modern human existence may have been a consequence of distinctive human capacities in social institution building.     

Shared Cultural History as a Predictor of Political and Economic Changes among Nation States

Political and economic risks arise from social phenomena that spread within and across countries. Regime changes, protest movements, and stock market and default shocks can have ramifications across the globe. Quantitative models have made great strides at predicting these events in recent decades but incorporate few explicitly measured cultural variables. However, in recent years cultural evolutionary theory has emerged as a major paradigm to understand the inheritance and diffusion of human cultural variation. Here, we combine these two strands of research by proposing that measures of socio-linguistic affiliation derived from language phylogenies track variation in cultural norms that influence how political and economic changes diffuse across the globe. First, we show that changes over time in a country’s democratic or autocratic character correlate with simultaneous changes among their socio-linguistic affiliations more than with changes of spatially proximate countries. Second, we find that models of changes in sovereign default status favor including socio-linguistic affiliations in addition to spatial data. These findings suggest that better measurement of cultural networks could be profoundly useful to policy makers who wish to diversify commercial, social, and other forms of investment across political and economic risks on an international scale.     

Evolutionary precursors of social norms in chimpanzees: a new approach

Moral behaviour, based on social norms, is commonly regarded as a hallmark of humans. Hitherto, humans are perceived to be the only species possessing social norms and to engage in moral behaviour. There is anecdotal evidence suggesting their presence in chimpanzees, but systematic studies are lacking. Here, we examine the evolution of human social norms and their underlying psychological mechanisms. For this, we distinguish between conventions, cultural social norms and universal social norms. We aim at exploring whether chimpanzees possess evolutionary precursors of universal social norms seen in humans. Chimpanzees exhibit important preconditions for their presence and enforcement: tolerant societies, well-developed social-cognitive skills and empathetic competence. Here, we develop a theoretical framework for recognizing different functional levels of social norms and distinguish them from mere statistical behavioural regularities. Quasi social norms are found where animals behave functionally moral without having moral emotions. In proto social norms, moral emotions might be present but cannot be collectivized due to the absence of a uniquely human psychological trait, i.e. shared intentionality. Human social norms, whether they are universal or cultural, involve moral emotions and are collectivized. We will discuss behaviours in chimpanzees that represent potential evolutionary precursors of human universal social norms, with special focus on social interactions involving infants. We argue that chimpanzee infants occupy a special status within their communities and propose that tolerance towards them might represent a proto social norm. Finally, we discuss possible ways to test this theoretical framework.     

Individual foraging specialisation in a social mammal: the European badger (Meles meles)

Individual specialisation has been identified in an increasing number of animal species and populations. However, in some groups, such as terrestrial mammals, it is difficult to disentangle individual niche variation from spatial variation in resource availability. In the present study, we investigate individual variation in the foraging niche of the European badger (Meles meles), a social carnivore that lives in a shared group territory, but forages predominantly alone. Using stable isotope analysis, we distinguish the extent to which foraging variation in badgers is determined by social and spatial constraints and by individual differences within groups. We found a tendency for individual badgers within groups to differ markedly and consistently in their isotope values, suggesting that individuals living with access to the same resources occupied distinctive foraging niches. Although sex had a significant effect on isotope values, substantial variation within groups occurred independently of age and sex. Individual differences were consistent over a period of several months and in some instances were highly consistent across the two years of the study, suggesting long-term individual foraging specialisations. Individual specialisation in foraging may, therefore, persist in populations of territorial species not solely as a result of spatial variation in resources, but also arising from individuals selecting differently from the same available resources. Although the exact cause of this behaviour is unknown, we suggest that specialisation may occur due to learning trade-offs which may limit individual niche widths. However, ecological factors at the group level, such as competition, may also influence the degree of specialisation.     

Natural cooperators: Food sharing in humans and other primates

The study of cooperation is rich with theoretical models and laboratory experiments that have greatly advanced our knowledge of human uniqueness, but have sometimes lacked ecological validity. We therefore emphasize the need to tie discussions of human cooperation to the natural history of our species and its closest relatives, focusing on behavioral contexts best suited to reveal underlying selection pressures and evolved decision rules. 1 - 3 Food sharing is a fundamental form of cooperation that is well‐studied across primates and is particularly noteworthy because of its central role in shaping evolved human life history, social organization, and cooperative psychology. 1 - 16 Here we synthesize available evidence on food sharing in humans and other primates, tracing the origins of offspring provisioning, mutualism, trade, and reciprocity throughout the primate order. While primates may gain some benefits from sharing, humans, faced with more collective action problems in a risky foraging niche, expanded on primate patterns to buffer risk and recruit mates and allies through reciprocity and signaling, and established co‐evolving social norms of production and sharing. Differences in the necessity for sharing are reflected in differences in sharing psychology across species, thus helping to explain unique aspects of our evolved cooperative psychology.     

Social learning in cooperative dilemmas

Helping is a cornerstone of social organization and commonplace in human societies. A major challenge for the evolutionary sciences is to explain how cooperation is maintained in large populations with high levels of migration, conditions under which cooperators can be exploited by selfish individuals. Cultural group selection models posit that such large-scale cooperation evolves via selection acting on populations among which behavioural variation is maintained by the cultural transmission of cooperative norms. These models assume that individuals acquire cooperative strategies via social learning. This assumption remains empirically untested. Here, I test this by investigating whether individuals employ conformist or payoff-biased learning in public goods games conducted in 14 villages of a forager–horticulturist society, the Pahari Korwa of India. Individuals did not show a clear tendency to conform or to be payoff-biased and are highly variable in their use of social learning. This variation is partly explained by both individual and village characteristics. The tendency to conform decreases and to be payoff-biased increases as the value of the modal contribution increases. These findings suggest that the use of social learning in cooperative dilemmas is contingent on individuals'' circumstances and environments, and question the existence of stably transmitted cultural norms of cooperation.     

Levels of selection, altruism, and primate behavior.

Altruistic behaviors seem anomalous from a traditional view of Darwinian natural selection, and evolutionary explanations for them have generated much discussion. The debate centers around four major explanations: classic individual-level selection, reciprocity and game theory, kin selection, and trait-group selection. The historical context and defining criteria of each model must be reviewed before its validity can be assessed. Of these proposed mechanisms, group selection historically has been the most controversial. Although the extent to which empirical data support group selection hypotheses is uncertain, there is evidence for group-level selection among avirulent virus strains and foraging ant queens. Researchers studying mammalian behavior, particularly primatologists, have largely dismissed models of group-level selection. Most discussion of altruism among primates has focused on differences in fitness among individuals within a single group, but students of altruistic behaviors exhibited by primates also need to investigate intergroup variation with respect to these behaviors. Various altruistic behaviors are likely to have evolved through different forms of selection, and each example of apparent altruism therefore needs to be evaluated separately.     

Review. Establishing an experimental science of culture: animal social diffusion experiments

A growing set of observational studies documenting putative cultural variations in wild animal populations has been complemented by experimental studies that can more rigorously distinguish between social and individual learning. However, these experiments typically examine only what one animal learns from another. Since the spread of culture is inherently a group-level phenomenon, greater validity can be achieved through 'diffusion experiments', in which founder behaviours are experimentally manipulated and their spread across multiple individuals tested. Here we review the existing corpus of 33 such studies in fishes, birds, rodents and primates and offer the first systematic analysis of the diversity of experimental designs that have arisen. We distinguish three main transmission designs and seven different experimental/control approaches, generating an array with 21 possible cells, 15 of which are currently represented by published studies. Most but not all of the adequately controlled diffusion experiments have provided robust evidence for cultural transmission in at least some taxa, with transmission spreading across populations of up to 24 individuals and along chains of up to 14 transmission events. We survey the achievements of this work, its prospects for the future and its relationship to diffusion studies with humans discussed in this theme issue and elsewhere.     

Gratitude Depends on the Relational Model of Communal Sharing

We studied the relation between benefits, perception of social relationships and gratitude. Across three studies, we provide evidence that benefits increase gratitude to the extent to which one applies a mental model of a communal relationship. In Study 1, the communal sharing relational model, and no other relational models, predicted the amount of gratitude participants felt after imagining receiving a benefit from a new acquaintance. In Study 2, participants recalled a large benefit they had received. Applying a communal sharing relational model increased feelings of gratitude for the benefit. In Study 3, we manipulated whether the participant or another person received a benefit from an unknown other. Again, we found that the extent of communal sharing perceived in the relationship with the stranger predicted gratitude. An additional finding of Study 2 was that communal sharing predicted future gratitude regarding the relational partner in a longitudinal design. To conclude, applying a communal sharing model predicts gratitude regarding concrete benefits and regarding the relational partner, presumably because one perceives the communal partner as motivated to meet one''s needs. Finally, in Study 3, we found in addition that being the recipient of a benefit without opportunity to repay directly increased communal sharing, and indirectly increased gratitude. These circumstances thus seem to favor the attribution of communal norms, leading to a communal sharing representation and in turn to gratitude. We discuss the importance of relational models as mental representations of relationships for feelings of gratitude.     

Pre-existing fairness concerns restrict the cultural evolution and generalization of inequitable norms in children

Many social exchanges produce benefits that would not exist otherwise, but anticipating conflicts about how to distribute these benefits can derail exchange and destroy the gains. Coordination norms can solve this problem by providing a shared understanding of how to distribute benefits, but such norms can also perpetuate group-level inequality. To examine how inequitable norms evolve culturally and whether they generalize from one setting to another, we conducted an incentivized lab-in-the-field experiment among kindergarten (5–6) and second-grade (8–9) children living in Switzerland (4′228 decisions collected from 326 children). In Part 1, we created two arbitrarily marked groups, triangles and circles. We randomly and repeatedly formed pairs with one triangle and one circle, and players in a pair played a simple bargaining game in which failure to agree destroyed the gains from social exchange. At the beginning of Part 1 we suggested a specific way to play the game. In symmetric treatments, this suggestion did not imply inequality between the groups, while in asymmetric treatments it did. Part 2 of the experiment addressed the generalization of norms. Retaining their group affiliations from Part 1, each child had to distribute resources between an in-group member and an out-group member. Children of both age groups in symmetric treatments used our suggestions about how to play the game to coordinate in Part 1. In asymmetric treatments, children followed our suggestions less consistently, which reduced coordination but moderated inequality. In Part 2, older children did not generalize privilege from Part 1. Rather, they compensated the underprivileged. Younger children neither generalized privilege nor compensated the underprivileged.     

Foraging behaviour alters with social environment in a juvenile songbird

Early independence from parents is a critical period where social information acquired vertically may become outdated, or conflict with new information. However, across natural populations, it is unclear if newly independent young persist in using information from parents, or if group-level effects of conformity override previous behaviours. Here, we test if wild juvenile hihi (Notiomystis cincta, a New Zealand passerine) retain a foraging behaviour from parents, or if they change in response to the behaviour of peers. We provided feeding stations to parents during chick-rearing to seed alternative access routes, and then tracked their offspring''s behaviour. Once independent, juveniles formed mixed-treatment social groups, where they did not retain preferences from their time with parents. Instead, juvenile groups converged over time to use one access route­ per group, and juveniles that moved between groups switched to copy the locally favoured option. Juvenile hihi did not copy specific individuals, even if they were more familiar with the preceding bird. Our study shows that early social experiences with parents affect initial foraging decisions, but social environments encountered later on can update transmission of arbitrary behaviours. This suggests that conformity may be widespread in animal groups, with potential cultural, ecological and evolutionary consequences.     

Within‐group relatedness is correlated with colony‐level social structure and reproductive sharing in a social fish

In group‐living species, the degree of relatedness among group members often governs the extent of reproductive sharing, cooperation and conflict within a group. Kinship among group members can be shaped by the presence and location of neighbouring groups, as these provide dispersal or mating opportunities that can dilute kinship among current group members. Here, we assessed how within‐group relatedness varies with the density and position of neighbouring social groups in Neolamprologus pulcher, a colonial and group‐living cichlid fish. We used restriction site‐associated DNA sequencing (RADseq) methods to generate thousands of polymorphic SNPs. Relative to microsatellite data, RADseq data provided much tighter confidence intervals around our relatedness estimates. These data allowed us to document novel patterns of relatedness in relation to colony‐level social structure. First, the density of neighbouring groups was negatively correlated with relatedness between subordinates and dominant females within a group, but no such patterns were observed between subordinates and dominant males. Second, subordinates at the colony edge were less related to dominant males in their group than subordinates in the colony centre, suggesting a shorter breeding tenure for dominant males at the colony edge. Finally, subordinates who were closely related to their same‐sex dominant were more likely to reproduce, supporting some restraint models of reproductive skew. Collectively, these results demonstrate that within‐group relatedness is influenced by the broader social context, and variation between groups in the degree of relatedness between dominants and subordinates can be explained by both patterns of reproductive sharing and the nature of the social landscape.