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.     

The evolutionary and ecological roots of human social organization

Social organization among human foragers is characterized by a three-generational system of resource provisioning within families, long-term pair-bonding between men and women, high levels of cooperation between kin and non-kin, and relatively egalitarian social relationships. In this paper, we suggest that these core features of human sociality result from the learning- and skill-intensive human foraging niche, which is distinguished by a late age-peak in caloric production, high complementarity between male and female inputs to offspring viability, high gains to cooperation in production and risk-reduction, and a lack of economically defensible resources. We present an explanatory framework for understanding variation in social organization across human societies, highlighting the interactive effects of four key ecological and economic variables: (i) the role of skill in resource production; (ii) the degree of complementarity in male and female inputs into production; (iii) economies of scale in cooperative production and competition; and (iv) the economic defensibility of physical inputs into production. Finally, we apply this framework to understanding variation in social and political organization across foraging, horticulturalist, pastoralist and agriculturalist societies.     

Multilevel modeling analysis of dyadic network data with an application to Ye'kwana food sharing

Behavioral ecologists have recently begun using multilevel modeling for the analysis of social behavior. We present a multilevel modeling formulation of the Social Relations Model that is well suited for the analysis of dyadic network data. This model, which we adapt for count data and small datasets, can be fitted using standard multilevel modeling software packages. We illustrate this model with an analysis of meal sharing among Ye'kwana horticulturalists in Venezuela. In this setting, meal sharing among households is predicted by an association index, which reflects the amount of time that members of the households are interacting. This result replicates recent findings that interhousehold food sharing is especially prevalent among households that interact and cooperate in multiple ways. We discuss opportunities for human behavioral ecologists to expand their focus to the multiple currencies and cooperative behaviors that characterize interpersonal relationships in preindustrial societies. We discuss possible extensions to this statistical modeling approach and applications to research by human behavioral ecologists and primatologists. Am J Phys Anthropol 157:507–512, 2015. © 2015 Wiley Periodicals, Inc.     

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.     

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.     

Tolerant food sharing and reciprocity is precluded by despotism among bonobos but not chimpanzees

Tolerant food sharing among human foragers can largely be explained by reciprocity. In contrast, food sharing among chimpanzees and bonobos may not always reflect reciprocity, which could be explained by different dominance styles: in egalitarian societies reciprocity is expressed freely, while in more despotic groups dominants may hinder reciprocity. We tested the degree of reciprocity and the influence of dominance on food sharing among chimpanzees and bonobos in two captive groups. First, we found that chimpanzees shared more frequently, more tolerantly, and more actively than bonobos. Second, among chimpanzees, food received was the best predictor of food shared, indicating reciprocal exchange, whereas among bonobos transfers were mostly unidirectional. Third, chimpanzees had a shallower and less linear dominance hierarchy, indicating that they were less despotic than bonobos. This suggests that the tolerant and reciprocal sharing found in chimpanzees, but not bonobos, was made possible by the absence of despotism. To investigate this further, we tested the relationship between despotism and reciprocity in grooming using data from an additional five groups and five different study periods on the main groups. The results showed that i) all chimpanzee groups were less despotic and groomed more reciprocally than bonobo groups, and ii) there was a general negative correlation between despotism and grooming reciprocity across species. This indicates that an egalitarian hierarchy may be more common in chimpanzees, at least in captivity, thus fostering reciprocal exchange. We conclude that a shallow dominance hierarchy was a necessary precondition for the evolution of human‐like reciprocal food sharing. Am J Phys Anthropol 143:41–51, 2010. © 2010 Wiley‐Liss, Inc.     

BaYaka forager food sharing networks in the Congo Basin: The roles of gender homophily and kin sharing

Objectives

Food sharing is a costly form of cooperation that was likely critical to human evolutionary success, including the emergence of human's life history strategy. Food sharing in human communities may be maintained through a number of pathways, including direct dyadic reciprocity, reputation-based processes, and kin-biased exchange. Differences in reproductive demands, labor, and cultural norms may also result in gendered differences in cooperative networks. Here, we examine cooperative networks in egalitarian BaYaka foragers from the Congo Basin.

Materials and Methods

We collected social network data from 112 adults in 41 households in this subsistence community. We implement a Bayesian latent network model to assess individual-, dyadic-, and block-level predictors of food sharing partners.

Results

Conditioning on covariates, we found limited evidence for direct dyadic reciprocity in food sharing. Despite local norms regarding prestige avoidance, we found status-based homophily. High-status individuals—council members and local healers—were more likely to share with one another. Importantly, our results highlight gender differences in patterns of food sharing, interacting with genetic relatedness. Women were more likely to share with one another, especially with kin as genetic relatedness increased.

Discussion

Our results align with evolutionary framing emphasizing kin selection in costly cooperation. The results showing that women cooperate with other women, particularly kin, also complement sex-based patterns in some other mammalian species, potentially reflecting the social support necessary to manage reproductive costs and childcare. BaYaka women's subsistence productivity and local cultural dynamics for autonomy and egalitarianism may likewise help facilitate women's preferential cooperation with one another.

     

Cooperation and conflict: The behavioral ecology of the sexual division of labor

When it comes to subsistence, men and women in almost all societies do it differently. One long-standing explanation for this sexual division of labor is that men and women pair up to provision offspring and specialize in subsistence activities in order to maximize household productivity. This model of cooperative parental provisioning has generally been supported by the proposal that both male and female reproductive success is maximized by provisioning current offspring rather than deserting them in order to seek new mating opportunities. But recent analyses of bird behavior have often failed to support this premise. We now know that among many species conflicting reproductive strategies between males and females often result in less than optimal compromises with regard to mating and parenting. This new focus on the role of sexual selection in creating compromise and conflict between the sexes has the potential to illuminate many puzzling aspects of human partnerships between men and women. To demonstrate its potential, I compare the explanatory power of a cooperative provisioning model of sex difference in human foraging and food sharing with a model incorporating conflicting reproductive goals.     

Cooperative breeding in South American hunter–gatherers

Evolutionary researchers have recently suggested that pre-modern human societies habitually practised cooperative breeding and that this feature helps explain human prosocial tendencies. Despite circumstantial evidence that post-reproductive females and extra-pair males both provide resources required for successful reproduction by mated pairs, no study has yet provided details about the flow of food resources by different age and sex categories to breeders and offspring, nor documented the ratio of helpers to breeders. Here, we show in two hunter–gatherer societies of South America that each breeding pair with dependent offspring on average obtained help from approximately 1.3 non-reproductive adults. Young married males and unmarried males of all ages were the main food providers, accounting for 93–100% of all excess food production available to breeding pairs and their offspring. Thus, each breeding pair with dependants was provisioned on average by 0.8 adult male helpers. The data provide no support for the hypothesis that post-reproductive females are the main provisioners of younger reproductive-aged kin in hunter–gatherer societies. Demographic and food acquisition data show that most breeding pairs can expect food deficits owing to foraging luck, health disabilities and accumulating dependency ratio of offspring in middle age, and that extra-pair provisioning may be essential to the evolved human life history.     

Bonobos Share with Strangers

Humans are thought to possess a unique proclivity to share with others – including strangers. This puzzling phenomenon has led many to suggest that sharing with strangers originates from human-unique language, social norms, warfare and/or cooperative breeding. However, bonobos, our closest living relative, are highly tolerant and, in the wild, are capable of having affiliative interactions with strangers. In four experiments, we therefore examined whether bonobos will voluntarily donate food to strangers. We show that bonobos will forego their own food for the benefit of interacting with a stranger. Their prosociality is in part driven by unselfish motivation, because bonobos will even help strangers acquire out-of-reach food when no desirable social interaction is possible. However, this prosociality has its limitations because bonobos will not donate food in their possession when a social interaction is not possible. These results indicate that other-regarding preferences toward strangers are not uniquely human. Moreover, language, social norms, warfare and cooperative breeding are unnecessary for the evolution of xenophilic sharing. Instead, we propose that prosociality toward strangers initially evolves due to selection for social tolerance, allowing the expansion of individual social networks. Human social norms and language may subsequently extend this ape-like social preference to the most costly contexts.     

Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees

Humans excel in cooperative exchanges between unrelated individuals. Although this trait is fundamental to the success of our species, its evolution and mechanisms are poorly understood. Other social mammals also build long-term cooperative relationships between non-kin, and recent evidence shows that oxytocin, a hormone involved in parent–offspring bonding, is likely to facilitate non-kin as well as kin bonds. In a population of wild chimpanzees, we measured urinary oxytocin levels following a rare cooperative event—food sharing. Subjects showed higher urinary oxytocin levels after single food-sharing events compared with other types of social feeding, irrespective of previous social bond levels. Also, urinary oxytocin levels following food sharing were higher than following grooming, another cooperative behaviour. Therefore, food sharing in chimpanzees may play a key role in social bonding under the influence of oxytocin. We propose that food-sharing events co-opt neurobiological mechanisms evolved to support mother–infant bonding during lactation bouts, and may act as facilitators of bonding and cooperation between unrelated individuals via the oxytocinergic system across social mammals.     

Showing off: Tests of an hypothesis about men's foraging goals

It is widely assumed that among hunter-gatherers, men work to provision their families. However, men may have more to gain by giving food to a wide range of companions who treat them favorably in return. If so, and if some resources better serve this end, men's foraging behavior should vary accordingly. Aspects of this hypothesis are tested on observations of food acquisition and sharing among Ache foragers of Eastern Paraguay. Previous analysis showed that different Ache food types were differently shared. Resources shared most widely were game animals. Further analysis and additional data presented here suggest a causal association between the wide sharing of game and the fact that men hunt and women do not. Data show that men preferentially target resources in both hunting and gathering which are more widely shared, resources more likely to be consumed outside their own nuclear families. These results have implications for 1) the identification of male reproductive trade-offs in human societies, 2) the view that families are units of common interest integrated by the sexual division of labor, 3) current reconstructions of the evolution of foraging and food sharing among early hominids, and 4) assessments of the role of risk and reciprocity in hunter-gatherer foraging strategies.     

Social structure and energy efficiency: A preliminary cross-national analysis

This study questions the assumption that there is an immutably positive relationship between per capita GNP and per capita energy consumption among human societies. A ratio of per capita GNP to per capita energy consumption ($U.S./kg coal equivalent) is proposed as a measure of energy efficiency for a cross-national analysis of 118 world nation-states and a subset of 25 developed market economies. This ratio is found to vary considerably, between 0.19 and 9.80. A review of literature suggests possible relationships among several sociodemographic characteristics of nations and levels of efficiency with which energy is converted into goods and services. Among the total sample, level of production (measured in terms of per capita GNP) bears a substantial inverse association with energy efficiency. When per capita GNP is held constant, agricultural share of gross domestic product and percentage of labor force in agriculture continue to be positively associated with energy efficiency among the total sample. Variables measuring defense expenditure, urbanization, and population density exhibit somewhat smaller multivariate relationships with energy efficiency when per capita GNP is controlled, i.e., these variables have significant multivariate parameters, but are less closely related to energy efficiency than level of production and agricultural composition of the economy and labor force. Agricultural composition of the economy and labor force is the major predictor of energy efficiency among the subset of 25 developed market economies. The results suggest that among the developed industrial societies level of production is less important than the composition of production activities in determining aggregate energy efficiency.This paper was originally presented at the annual meeting of the North Central Sociological Association, Pittsburgh, Pennsylvania, May 1977. This research was supported by funds from the Michigan Agricultural Experiment Station, the Ohio Agricultural Research and Development Center, and the National Research Council, National Academy of Sciences.     

The Organization of Male and Female Labor in Foraging Societies: Implications for Early Paleoindian Archaeology

I use cross-cultural ethnographic data to explore the relationship between male and female subsistence labor among hunter-gatherer populations by examining data regarding resource procurement, time allocation, and task differentiation between the sexes relative to dependence on hunted foods. The findings indicate that female foragers generally perform a variety of nonsubsistence collection activities and preferentially procure high-return resources in hunting-based economies. I develop ideas about predictable relationships concerning the amount of time female foragers expend on subsistence and technological tasks relative to the dietary contribution of meat. I then use ethnographic trends to evaluate archaeological assumptions regarding the sexual division of labor in prehistoric foraging contexts, focusing on the dichotomous views of Clovis labor organization. I argue that archaeological interpretations of prehistoric labor roles in hunting-based foraging societies are commonly polarized between stereotypical views of male and female subsistence behaviors. I develop an interpretation of Early Paleoindian labor organization, emphasizing female labor in the production of material goods and the procurement of low-risk plant and animal resources based on global economic trends among foragers.     

“It's a Wonderful Life”: signaling generosity among the Ache of Paraguay

Intensive food sharing among foragers and horticulturists is commonly explained as a means of reducing the risk of daily shortfalls, ensuring adequate daily consumption for all group members who actively pool resources. Consistently high food producers who give more than they receive, however, gain the least risk-reduction benefit from this daily pooling because they are the least likely to go without food on any given day. Why then do some high producers consistently share food, and why do some average producers share proportionally more food than others? We propose that although these individuals may not receive the same amounts they give (i.e., strict Tit-for-Tat), one explanation for their generosity is that they receive additional food during hard times. These include brief episodes of sickness, disease, injury, or accidents—fairly common events in traditional societies that can render individuals incapable of producing food, thereby having long-term effects on morbidity and fecundity and ultimately on lifetime reproductive success. Data collected among the Ache, a group of South American forager-horticulturists, indicate that those who shared and produced more than average (signaling cooperative intent and/or ability to produce) were rewarded with more food from more people when injured or sick than those who shared and produced below average. These results, framed within the context of tradeoffs between short-term and long-term fitness, may provide insight into motivations behind costly expenditures for establishing and reinforcing status and reputation.     

Forum: The eusociality continuum

Eusocial societies are traditionally characterized by a reproductivedivision of labor, an overlap of generations, and cooperativecare of the breeders' young. Eusociality was once thought tooccur only in termites, ants, and some bee and wasp species,but striking evolutionary convergences have recently becomeapparent between the societies of these insects and those ofcooperatively breeding birds and mammals. These parallels haveblurred distinctions between cooperative breeding and eusociality,leading to calls for either drastically restricting or expandingusage of these terms. We favor the latter approach. Cooperativebreeding and eusociality are not discrete phenomena, but ratherform a continuum of fundamentally similar social systems whosemain differences lie in the distribution of lifetime reproductivesuccess among group members. Therefore we propose to array vertebrateand invertebrate cooperative breeders along a common axis, representinga standardized measure of reproductive variance, and to dropsuch (loaded) terms as "primitive" and "advanced" eusociality.The terminology we propose unites all occurrences of alloparentalhelping of kin under a single theoretical umbrella (e.g., Hamilton'srule). Thus, cooperatively breeding vertebrates can be regardedas eusocial, just as eusocial invertebrates are cooperativebreeders. We believe this integrated approach will foster potentiallyrevealing cross-taxon comparisons, which are essential to understandingsocial evolution in birds, mammals, and insects.     

The Insectan Apes

I present evidence that humans have evolved convergently to social insects with regard to a large suite of social, ecological, and reproductive phenotypes. Convergences between humans and social insects include: (1) groups with genetically and environmentally defined structures; (2) extensive divisions of labor; (3) specialization of a relatively restricted set of females for reproduction, with enhanced fertility; (4) extensive extramaternal care; (5) within-group food sharing; (6) generalized diets composed of high-nutrient-density food; (7) solicitous juveniles, but high rates of infanticide; (8) ecological dominance; (9) enhanced colonizing abilities; and (10) collective, cooperative decision-making. Most of these convergent phenotypic adaptations stem from reorganization of key life-history trade-offs due to behavioral, physiological, and life-historical specializations. Despite their extensive socioreproductive overlap with social insects, humans differ with regard to the central aspect of eusociality: reproductive division of labor. This difference may be underpinned by the high energetic costs of producing offspring with large brains.     

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 Archaeology of Food Preference

Food preference is a socially constructed concept in which both consumers and producers define what is "good to eat." Staple crops and daily meals are an important component of these definitions, as the regular use of particular foods reinforces norms of identity. Food preferences also affect agricultural systems because choices among cultivars are based on social needs in addition to economic variables such as yield and caloric value. Through textual and archaeological evidence, the trajectory of rice production is examined for Sri Lanka, the Brahmaputra Valley, the Tamil region, and Vijayanagara. In these regions and elsewhere in South Asia, shared ideologies of food preference resulted in a consensus mode of agricultural production: Irrigation works increased the tax base for political leaders and the donation base for temple economies, but they also benefited local inhabitants who would have been able to partake of a preferred food on a more regular basis.     

The evolution of human parental care and recruitment of juvenile help

Provisioning of juveniles is a defining characteristic of human life history. Human children are also unusual in cooperating with their siblings, mothers and other adults in the exchange of resources and labor. This article highlights this distinctly human and twofold nature of juvenility within the context of life history evolution and cooperative breeding. Juveniles benefit from continued investment and from helping to support their siblings during a life stage when they cannot contribute to their own reproduction. Rather than juvenile dependence signifying a costly extension of parental care, juvenile provisioning and help are suggested to develop in tandem with the broader pattern of food sharing and division of labor that characterizes human subsistence and sociality.