Women's subsistence strategies predict fertility across cultures, but context matters.

While it is commonly assumed that farmers have higher, and foragers lower, fertility compared to populations practicing other forms of subsistence, robust supportive evidence is lacking. We tested whether subsistence activities-incorporating market integration-are associated with fertility in 10,250 women from 27 small-scale societies and found considerable variation in fertility. This variation did not align with group-level subsistence typologies. Societies labeled as "farmers" did not have higher fertility than others, while "foragers" did not have lower fertility. However, at the individual level, we found strong evidence that fertility was positively associated with farming and moderate evidence of a negative relationship between foraging and fertility. Markers of market integration were strongly negatively correlated with fertility. Despite strong cross-cultural evidence, these relationships were not consistent in all populations, highlighting the importance of the socioecological context, which likely influences the diverse mechanisms driving the relationship between fertility and subsistence.

Hormone physiology and sleep dynamics among BaYaka foragers of the Congo Basin: Gendered associations between nighttime activity, testosterone, and cortisol.

Sleep quality is an important contributor to health disparities and affects the physiological function of the immune and endocrine systems, shaping how resources are allocated to life history demands. Past work in industrial and post-industrial societies has shown that lower total sleep time (TST) or more disrupted nighttime sleep are linked to flatter diurnal slopes for cortisol and lower testosterone production. There has been little focus on these physiological links in other socio-ecological settings where routine sleep conditions and nighttime activity demands differ. We collected salivary hormone (testosterone, cortisol) and actigraphy-based sleep data from Congolese BaYaka foragers (N = 39), who have relatively short and fragmented nighttime sleep, on average, in part due to their typical social sleep conditions and nighttime activity. The hormone and sleep data collections were separated by an average of 11.23 days (testosterone) and 2.84 days (cortisol). We found gendered links between nighttime activity and adults' hormone profiles. Contrary to past findings in Euro-American contexts, BaYaka men who were more active at night, on average, had higher evening testosterone than those with lower nighttime activity, with a relatively flat slope relating nighttime activity and evening testosterone in women. Women had steeper diurnal cortisol curves with less disrupted sleep. Men had steeper cortisol diurnal curves if they were more active at night. BaYaka men often hunt and socialize when active at night, which may help explain these patterns. Overall, our findings indicate that the nature of nighttime activities, including their possible social and subsistence contexts, are potentially important modifiers of sleep quality-physiology links, meriting further research across contexts.

Girls in early childhood increase food returns of nursing women during subsistence activities of the BaYaka in the Republic of Congo.

Nursing mothers face an energetic trade-off between infant care and work. Under pooled energy budgets, this trade-off can be reduced by assistance in food acquisition and infant care tasks from non-maternal carers. Across cultures, children also often provide infant care. Yet the question of who helps nursing mothers during foraging has been understudied, especially the role of children. Using focal follow data from 140 subsistence expeditions by BaYaka women in the Republic of Congo, we investigated how potential support from carers increased mothers' foraging productivity. We found that the number of girls in early childhood (ages 4-7 years) in subsistence groups increased food returns of nursing women with infants (kcal collected per minute). This effect was stronger than that of other adult women, and older girls in middle childhood (ages 8-13 years) and adolescence (ages 14-19 years). Child helpers were not necessarily genetically related to nursing women. Our results suggest that it is young girls who provide infant care while nursing mothers are acquiring food-by holding, monitoring and playing with infants-and, thus, that they also contribute to the energy pool of the community during women's subsistence activities. Our study highlights the critical role of children as carers from early childhood.

Gender-Typed and Gender-Segregated Play Among Tanzanian Hadza and Congolese BaYaka Hunter-Gatherer Children and Adolescents.

Few data exist on gender-typed and gender-segregated play in hunter-gatherer societies, despite their unique demographic and cultural features which may influence children's gendered play. Using naturalistic observations of Hadza (N = 46, 41% female) and BaYaka (N = 65, 48% female) hunter-gatherer 3- to 18-year-olds from Tanzania and the Republic of Congo, we showed that access to playmates was negatively associated with playing in mixed-gender groups. Young boys did not engage in more rough-and-tumble play than girls, but adolescent boys participated in this type of play more than adolescent girls. Children were also more likely to participate in work-themed play which conformed to gender norms within their society. Findings are discussed within the context of gendered division of labor, child autonomy, and demography.

Sex differences in daily activity intensity and energy expenditure and their relationship to cortisol among BaYaka foragers from the Congo Basin.

OBJECTIVES: The pooling of energetic resources and food sharing have been widely documented among hunter-gatherer societies. Much less is known about how the energetic costs of daily activities are distributed across individuals in such groups, including between women and men. Moreover, the metabolic physiological correlates of those activities and costs are relatively understudied. MATERIALS AND METHODS: Here, we tracked physical activity, energy expenditure (EE), and cortisol production among Congo Basin BaYaka foragers engaged in a variety of daily subsistence activities (n = 37). Given its role in energy mobilization, we measured overall daily cortisol production and short-term cortisol reactivity through saliva sampling; we measured physical activity levels and total EE via the wGT3X-bt actigraph and heart rate monitor. RESULTS: We found that there were no sex differences in likelihood of working in common activity locations (forest, garden, house). Across the day, women spent greater percentage time in moderate-to-vigorous physical activity (%MVPA) and had lower total EE than men. Females with higher EE (kCal/hr) produced greater cortisol throughout the day. Though not statistically significant, we also found that individuals with greater %MVPA had larger decreases in cortisol reactivity. DISCUSSION: BaYaka women sustained higher levels of physical activity but incurred lower energetic costs than men, even after factoring in sex differences in body composition. Our findings suggest that the distribution of physical activity demands and costs are relevant to discussions regarding how labor is divided and community energy budgets take shape in such settings.

Fatherhood, egalitarianism, and child health in two small-scale societies in the Republic of the Congo.

OBJECTIVES: The study goals were to (a) characterize the cultural model of fatherhood among the BaYaka, a community of egalitarian foragers in the Republic of the Congo; (b) test if BaYaka fathers' quality in relation to the cultural model predicts their children's energetic status; and (c) compare the variance in BaYaka children's energetic status to that of children of neighboring Bondongo fisher-farmers, among whom there is less cooperative caregiving, less resource sharing, and greater social inequality. METHODS: We used informal interviews to establish the cultural model of fatherhood, which we used to build a peer ranking task to quantify father quality. Children's energetic status was assessed by measuring height, weight, and triceps skinfold thickness. We then tested for associations between father quality scores derived from the ranking task and children's energetic status using ordinary least squares regression. Equality of variance tests were used to compare BaYaka and Bondongo children's energetic statuses. RESULTS: The BaYaka described fathers as responsible for acquiring resources and maintaining marital harmony, welcoming others to the community and sharing well with them, and teaching their children about the forest. Agreement on men's quality in these domains was high, but father quality did not significantly predict children's energetic status. BaYaka children had lower variance in energetic status overall compared to Bondongo children. CONCLUSIONS: We suggest that the core BaYaka values and practices that maintain egalitarian social relations and distribution of resources help buffer children's health and well-being from variation in their fathers' qualities in culturally valued domains.

Epigenetic aging in children from a small-scale farming society in The Congo Basin: Associations with child growth and family conflict.

Developmental environments influence individuals' long-term health trajectories, and there is increasing emphasis on understanding the biological pathways through which this occurs. Epigenetic aging evaluates DNA methylation at a suite of distinct CpG sites in the genome, and epigenetic age acceleration (EAA) is linked to heightened chronic morbidity and mortality risks in adults. Consequently, EAA provides insights on trajectories of biological aging, which early life experiences may help shape. However, few studies have measured correlates of children's epigenetic aging, especially outside of the U.S. and Europe. In particular, little is known about how children's growth and development relate to EAA in ecologies in which energetic and pathogenic stressors are commonplace. We studied EAA from dried blood spots among Bondongo children (n = 54) residing in a small-scale, fisher-farmer society in a remote region of the Republic of the Congo. Here, infectious disease burdens and their resultant energy demands are high. Children who were heavier for height or taller for age, respectively, exhibited greater EAA, including intrinsic EAA, which is considered to measure EAA internal to cells. Furthermore, we found that children in families with more conflict between parents had greater intrinsic EAA. These results suggest that in contexts in which limited energy must be allocated to competing demands, more investment in growth may coincide with greater EAA, which parallels findings in European children who do not face similar energetic constraints. Our findings also indicate that associations between adverse family environments and greater intrinsic EAA were nonetheless observable but only after adjustment for covariates relevant to the energetically and immunologically demanding nature of the local ecology.

Testosterone, fathers as providers and caregivers, and child health: Evidence from fisher-farmers in the Republic of the Congo.

Males in vertebrate species with biparental care commonly face a life history trade-off between investing in mating versus parenting effort. Among these males, testosterone is frequently elevated during mating and competition and reduced when males help raise offspring. These physiological patterns may be adaptive, increasing males' fitness through investments in young. However, for some species, including humans, indirect parenting often benefits young but can also involve male competition and risk-taking behavior and may be facilitated by elevated testosterone. Despite potential adaptive functions of biological responses to invested fatherhood, few if any mammalian studies have linked fathers' testosterone to offspring outcomes; no studies in humans have. Using data from a small-scale society of fisher-farmers from the Republic of the Congo, we find that fathers who were rated as better providers by their peers had higher testosterone, compared to other fathers in their community. However, children whose fathers had middle-range T compared to fathers with higher or lower levels had better energetic status (higher BMI; greater triceps skinfold thickness). Fathers' indirect and direct care helped to account for these associations between paternal T and children's energetic profiles. Given that human paternal direct and, especially, indirect care are thought to have been important evolutionarily and remain so in many contemporary societies, these findings help to shed light on the facultative nature of human biological responses to fatherhood and the relevance of these factors to children's well-being.

Human preferences for sexually dimorphic faces may be evolutionarily novel.

A large literature proposes that preferences for exaggerated sex typicality in human faces (masculinity/femininity) reflect a long evolutionary history of sexual and social selection. This proposal implies that dimorphism was important to judgments of attractiveness and personality in ancestral environments. It is difficult to evaluate, however, because most available data come from large-scale, industrialized, urban populations. Here, we report the results for 12 populations with very diverse levels of economic development. Surprisingly, preferences for exaggerated sex-specific traits are only found in the novel, highly developed environments. Similarly, perceptions that masculine males look aggressive increase strongly with development and, specifically, urbanization. These data challenge the hypothesis that facial dimorphism was an important ancestral signal of heritable mate value. One possibility is that highly developed environments provide novel opportunities to discern relationships between facial traits and behavior by exposing individuals to large numbers of unfamiliar faces, revealing patterns too subtle to detect with smaller samples.

Children's Play and Culture Learning in an Egalitarian Foraging Society.

Few systematic studies of play in foragers exist despite their significance for understanding the breadth of contexts for human development and the ontogeny of cultural learning. Forager societies lack complex social hierarchies, avenues for prestige or wealth accumulation, and formal educational institutions, and thereby represent a contrast to the contexts of most play research. Analysis of systematic observations of children's play among Aka forest foragers (n = 50, ages 4-16, M = 9.5) and Ngandu subsistence farmers (n = 48, ages 4-16, M = 9.1) collected in 2010 illustrates that while play and work trade off during development in both groups, and consistent patterns in sex-role development are evident, Aka children engage in significantly less rough-and-tumble play and competitive games than children among their socially stratified farming neighbors.

Ontogeny of prosocial behavior across diverse societies.

Humans are an exceptionally cooperative species, but there is substantial variation in the extent of cooperation across societies. Understanding the sources of this variability may provide insights about the forces that sustain cooperation. We examined the ontogeny of prosocial behavior by studying 326 children 3-14 y of age and 120 adults from six societies (age distributions varied across societies). These six societies span a wide range of extant human variation in culture, geography, and subsistence strategies, including foragers, herders, horticulturalists, and urban dwellers across the Americas, Oceania, and Africa. When delivering benefits to others was personally costly, rates of prosocial behavior dropped across all six societies as children approached middle childhood and then rates of prosociality diverged as children tracked toward the behavior of adults in their own societies. When prosocial acts did not require personal sacrifice, prosocial responses increased steadily as children matured with little variation in behavior across societies. Our results are consistent with theories emphasizing the importance of acquired cultural norms in shaping costly forms of cooperation and creating cross-cultural diversity.

Women's subsistence networks scaffold cultural transmission among BaYaka foragers in the Congo Basin.

In hunter-gatherer societies, women's subsistence activities are crucial for food provisioning and children's social learning but are understudied relative to men's activities. To understand the structure of women's foraging networks, we present 230 days of focal-follow data in a BaYaka community. To analyze these data, we develop a stochastic blockmodel for repeat observations with uneven sampling. We find that women's subsistence networks are characterized by cooperation between kin, gender homophily, and mixed age-group composition. During early childhood, individuals preferentially coforage with adult kin, but those in middle childhood and adolescence are likely to coforage with nonkin peers, providing opportunities for horizontal learning. By quantifying the probability of coforaging ties across age classes and relatedness levels, our findings provide insights into the scope for social learning during women's subsistence activities in a real-world foraging population and provide ground-truth values for key parameters used in formal models of cumulative culture.

Dyadic inter-group cooperation in shotgun hunting activities in a Congo Basin village.

Understanding the dynamics of inter-group cooperation in human adaptation has been the subject of recent empirical and theoretical studies in evolutionary anthropology, beginning to fill gaps in our knowledge of how interactions across political, economic and social domains can - and often do - lead to stable, large-scale cooperation. Here we investigate dyadic intergroup cooperation in shotgun hunting in the Republic of the Congo. In the Congo Basin, inter-group cooperation between foragers and farmers is at the centre of an exchange system maintained by traditional norms and institutions such as fictive kinship. Here, we focused on what factors predict cooperative shotgun hunting exchanges between BaYaka and Yambe. We conducted structured interviews with 48 BaYaka hunters and 18 Yambe men who organise hunts in a village along the Motaba River. We used Bayesian multilevel regression models to investigate the influence of Yambe and BaYaka attributes on probability of dyadic cooperation. We found that BaYaka men's reputations as skilled hunters and their family size each predicted cooperation in shotgun hunting, whereas there was no effect of Yambe attributes (status, wealth, family size). We discuss the results in terms of evolutionary models of men as hunters and inter-group cooperation, as well as biodiversity conservation implications.

Intergroup Cooperation in Shotgun Hunting Among BaYaka Foragers and Yambe Farmers from the Republic of the Congo.

Whereas many evolutionary models emphasize within-group cooperation or between-group competition in explaining human large-scale cooperation, recent work highlights a critical role for intergroup cooperation in human adaptation. Here we investigate intergroup cooperation in the domain of shotgun hunting in northern Republic of the Congo. In the Congo Basin broadly, forest foragers maintain relationships with neighboring farmers based on systems of exchange regulated by norms and institutions such as fictive kinship. In this study, we examine how relationships between Yambe farmers and BaYaka foragers support stable intergroup cooperation in the domain of shotgun hunting. In the study village, shotgun hunting is based on a specialization-based exchange wherein Yambe farmers contribute shotguns and access to markets to buy cartridges and sell meat while BaYaka foragers contribute their specialized forest knowledge and skill. To understand how costs and benefits are distributed, we conducted structured interviews with 77 BaYaka hunters and 15 Yambe gun owners and accompanied hunters on nine hunting trips. We found that hunts are organized in a conventional manner within a fictive kinship structure, consistent with the presence of intercultural mechanisms to stabilize cooperation. However, because bushmeat demand is high, gun owners can gain significant cash profit, while compensating hunters only with cigarettes, alcohol, and a traditional hunter's portion of meat. To level payoffs, hunters strategically hide kills or cartridges from gun owners to feed their own families. Our results illustrate how each group prioritizes different currencies (e.g., cash, meat, family, intergroup relations) and provide insights into how intergroup cooperation is stabilized in this setting. The example of this long-standing intergroup cooperative system is discussed in terms of its contemporary entwinement with logging, the bushmeat trade, and growing market intersection.

Anointing variation across wild capuchin populations: a review of material preferences, bout frequency and anointing sociality in Cebus and Sapajus.

The frequency of anointing bouts and the materials used for self- and social anointing vary across capuchin species in captivity, but there is little published data on capuchin anointing in the wild. Here we present previously unpublished data on anointing behaviors from capuchin monkey populations at ten different field sites and incorporate these data into a review of the anointing literature for captive and wild capuchins. Using a comparative phylogenetic framework, we test four hypotheses derived primarily from captive literature for variation in anointing between wild untufted capuchins (Cebus) and tufted capuchins (Sapajus), including that (1) the frequency of anointing is higher in Cebus, (2) Cebus uses a higher proportion of plant species to insect species for anointing compared with Sapajus, (3) anointing material diversity is higher in Cebus, and (4) social indices of anointing are higher in Cebus. We found that wild Cebus anoints more with plant parts, including fruits, whereas wild Sapajus anoints more with ants and other arthropods. Cebus capucinus in particular uses more plant species per site for anointing compared with other capuchins and may specialize in anointing as an activity independent from foraging, whereas most other capuchin species tend to eat the substances they use for anointing. In agreement with captive studies, we found evidence that wild Cebus anoints at a significantly higher frequency than Sapajus. However, contrary to the captive literature, we found no difference in the range of sociality for anointing between Cebus and Sapajus in the wild. We review anointing in the context of other Neotropical primate rubbing behaviors and consider the evidence for anointing as self-medication; as a mechanism for enhanced sociality; and as a behavioral response to chemical stimuli.

Social ties in the Congo Basin: insights into tropical forest adaptation from BaYaka and their neighbours.

Investigating past and present human adaptation to the Congo Basin tropical forest can shed light on how climate and ecosystem variability have shaped human evolution. Here, we first review and synthesize genetic, palaeoclimatological, linguistic and historical data on the peopling of the Congo Basin. While forest fragmentation led to the increased genetic and geographical divergence of forest foragers, these groups maintained long-distance connectivity. The eventual expansion of Bantu speakers into the Congo Basin provided new opportunities for forging inter-group links, as evidenced by linguistic shifts and historical accounts. Building from our ethnographic work in the northern Republic of the Congo, we show how these inter-group links between forest forager communities as well as trade relationships with neighbouring farmers facilitate adaptation to ecoregions through knowledge exchange. While researchers tend to emphasize forager-farmer interactions that began in the Iron Age, we argue that foragers' cultivation of relational wealth with groups across the region played a major role in the initial occupation of the Congo Basin and, consequently, in cultural evolution among the ancestors of contemporary peoples. This article is part of the theme issue 'Tropical forests in the deep human past'.

The social learning and development of intra- and inter-ethnic sharing norms in the Congo Basin: A registered report protocol.

Compared to other species, the extent of human cooperation is unparalleled. Such cooperation is coordinated between community members via social norms. Developmental research has demonstrated that very young children are sensitive to social norms, and that social norms are internalized by middle childhood. Most research on social norm acquisition has focused on norms that modulated intra-group cooperation. Yet around the world, multi-ethnic communities also cooperate, and this cooperation is often shaped by distinct inter-group social norms. In the present study, we will investigate whether inter-ethnic and intra-ethnic social norm acquisition follows the same, or distinct, developmental trajectories. Specifically, we will work with BaYaka foragers and Bandongo fisher-farmers who inhabit multi-ethnic villages in the Republic of the Congo. In these villages, inter-ethnic cooperation is regulated by sharing norms. Through interviews with adult participants, we will provide the first descriptive account of the timing and mechanism by which BaYaka and Bandongo learn to share with out-group members. Children (5-17 years) and adults (17+ years) will also participate in a modified Dictator Game to investigate the developmental trajectories of children's intra- and inter-ethnic sharing choices. Based on our ethnographic knowledge of the participating communities, we predict that children's intra-ethnic sharing choices in the Dictator Game will match those of adults at an earlier age than their inter-ethnic sharing choices. We will analyze our data using logistic Bayesian modelling.

Links between household and family social dynamics with sleep profiles among BaYaka foragers of the Congo Basin.

Given the contributions of sleep to a range of health outcomes, there is substantial interest in ecological and environmental factors, including psychosocial contexts, that shape variation in sleep between individuals and populations. However, the links between social dynamics and sleep are not well-characterized beyond Euro-American settings, representing a gap in knowledge regarding the way that local socio-ecological conditions interrelate with sleep profiles across diverse settings. Here, we focused on data from a subsistence-level society in Republic of the Congo to test for links between the household/family social environment and sleep measures. Specifically, we used actigraphy-derived sleep data (N = 49; 318 nights) from two community locations (a village and rainforest camp) among BaYaka foragers, who are members of a remote, non-industrialized subsistence society in the Congo Basin. We drew on social dynamics that have been previously linked to sleep variation in Euro-American contexts, including: household crowding, same surface cosleeping, and marital conflict. We examined the following sleep measures: total sleep time (TST), total 24-h sleep time (TTST), and sleep quality (fragmentation). BaYaka adults had shorter and lower quality sleep when their shared sleeping space was more crowded. In the village, parents with breastfeeding-aged infants had longer TTST and higher quality sleep than adults without infants, contrasting with results from other cultural contexts. Based on peer rankings of marital conflict, husbands showed longer and higher quality sleep in less conflicted marriages. Women showed the opposite pattern. These counter-intuitive findings for women may reflect the limitations of the measurement for wives' marital experiences. In total, these results point to the importance of considering local socio-ecological conditions to sleep profiles and underscore the need for expanded study of sleep and health outcomes in settings where shared sleep in constrained space is routine practice.

Socioecology shapes child and adolescent time allocation in twelve hunter-gatherer and mixed-subsistence forager societies.

A key issue distinguishing prominent evolutionary models of human life history is whether prolonged childhood evolved to facilitate learning in a skill- and strength-intensive foraging niche requiring high levels of cooperation. Considering the diversity of environments humans inhabit, children's activities should also reflect local social and ecological opportunities and constraints. To better understand our species' developmental plasticity, the present paper compiled a time allocation dataset for children and adolescents from twelve hunter-gatherer and mixed-subsistence forager societies (n = 690; 3-18 years; 52% girls). We investigated how environmental factors, local ecological risk, and men and women's relative energetic contributions were associated with cross-cultural variation in child and adolescent time allocation to childcare, food production, domestic work, and play. Annual precipitation, annual mean temperature, and net primary productivity were not strongly associated with child and adolescent activity budgets. Increased risk of encounters with dangerous animals and dehydration negatively predicted time allocation to childcare and domestic work, but not food production. Gender differences in child and adolescent activity budgets were stronger in societies where men made greater direct contributions to food production than women. We interpret these findings as suggesting that children and their caregivers adjust their activities to facilitate the early acquisition of knowledge which helps children safely cooperate with adults in a range of social and ecological environments. These findings compel us to consider how childhood may have also evolved to facilitate flexible participation in productive activities in early life.

Children's fingernail cortisol among BaYaka foragers of the Congo Basin: associations with fathers' roles.

Children and mothers' cortisol production in response to family psychosocial conditions, including parenting demands, family resource availability and parental conflict, has been extensively studied in the United States and Europe. Less is known about how such family dynamics relate to family members' cortisol in societies with a strong cultural emphasis on cooperative caregiving. We studied a cumulative indicator of cortisol production, measured from fingernails, among BaYaka forager children (77 samples, n = 48 individuals) and their parents (78 samples, n = 49) in the Congo Basin. Men ranked one another according to locally valued roles for fathers, including providing resources for the family, sharing resources in the community and engaging in less marital conflict. Children had higher cortisol if their parents were ranked as having greater parental conflict, and their fathers were seen as less effective providers and less generous sharers of resources in the community. Children with lower triceps skinfold thickness (an indicator of energetic condition) also had higher cortisol. Parental cortisol was not significantly correlated to men's fathering rankings, including parental conflict. Our results indicate that even in a society in which caregiving is highly cooperative, children's cortisol production was nonetheless correlated to parental conflict as well as variation in locally defined fathering quality. This article is part of the theme issue 'Multidisciplinary perspectives on social support and maternal-child health'.