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.

Reproductive inequality in humans and other mammals.

To address claims of human exceptionalism, we determine where humans fit within the greater mammalian distribution of reproductive inequality. We show that humans exhibit lower reproductive skew (i.e., inequality in the number of surviving offspring) among males and smaller sex differences in reproductive skew than most other mammals, while nevertheless falling within the mammalian range. Additionally, female reproductive skew is higher in polygynous human populations than in polygynous nonhumans mammals on average. This patterning of skew can be attributed in part to the prevalence of monogamy in humans compared to the predominance of polygyny in nonhuman mammals, to the limited degree of polygyny in the human societies that practice it, and to the importance of unequally held rival resources to women's fitness. The muted reproductive inequality observed in humans appears to be linked to several unusual characteristics of our species-including high levels of cooperation among males, high dependence on unequally held rival resources, complementarities between maternal and paternal investment, as well as social and legal institutions that enforce monogamous norms.

Post-marital residence patterns and the timing of reproduction: evidence from a matrilineal society.

Humans exhibit a broad range of post-marital residence patterns and there is growing recognition that post-marital residence predicts women's reproductive success; however, the nature of the relationship is probably dependent on whether co-resident kin are cooperators or competitors. Here, we explore this relationship in a Tibetan population, where couples practice a mixture of post-marital residence patterns, co-residing in the same village with the wife's parents, the husband's parents or endogamously with both sets of parents. Using detailed demographic data from 17 villages we find that women who live with only their own parents have an earlier age at first birth (AFB) and age at last birth (ALB) than women who live with only their parents-in-law. Women who co-reside with both sets of parents have the earliest AFB and ALB. However, those with co-resident older siblings postponed reproduction, suggestive of competition-related delay. Shifts to earlier reproductive timing were also observed in relation to the imposition of family planning policies, in line with Fisherian expectations. Our study provides evidence of the costs and benefits to women's direct fitness of co-residing with different kin, against a backdrop of adaptive responses to cultural constraints on completed fertility.

Testing the buffering hypothesis: Breastfeeding problems, cessation, and social support in the UK.

OBJECTIVES: Physical breastfeeding problems can lead women to terminate breastfeeding earlier than planned. In high-income countries such as the UK, breastfeeding problems have been attributed to the cultural and individual "inexperience" of breastfeeding, ultimately leading to lower breastfeeding rates. Yet, cross-cultural evidence suggests breastfeeding problems still occur in contexts where breastfeeding is common, prolonged, and seen publicly. This suggests breastfeeding problems are not unusual and do not necessarily lead to breastfeeding cessation. As humans evolved to raise children cooperatively, what matters for breastfeeding continuation may be the availability of social support during the postnatal period. Here, we test the hypothesis that social support buffers mothers from the negative impact breastfeeding problems have on duration. METHODS: We run Cox models on a sample of 565 UK mothers who completed a retrospective online survey about infant feeding and social support in 2017-2018. RESULTS: Breastfeeding problems were important predictors of cessation; however, the direction of the effect was dependent on the problem type and type of support from a range of supporters. Helpful support for discomfort issues (blocked ducts, too much milk) was significantly associated with reduced hazards of cessation, as predicted. However, helpful support for reported milk insufficiency was assoicated with an increased hazard of cessation. CONCLUSIONS: Experiencing breastfeeding problems is the norm, but its impact may be mitigated via social support. Working from an interdisciplinary approach, our results highlight that a wide range of supporters who provide different types of support have potential to influence maternal breastfeeding experience.

Reconstructing prehistoric demography: What role for extant hunter-gatherers?

Demography is central to biological, behavioral, and cultural evolution. Knowledge of the demography of prehistoric populations of both Homo sapiens and earlier members of the genus Homo is, therefore, key to the study of human evolution. Unfortunately, demographic processes (fertility, mortality, migration) leave little mark on the archeological and paleoanthropological records. One common solution to this issue is the application of demographic data from extant hunter-gatherers to prehistory. With the aim of strengthening this line of enquiry, here we outline some pitfalls and their interpretative implications. In doing so, we provide recommendations about the application of hunter-gatherer data to the study of demographic trends throughout human evolution. We use published demographic data from extant hunter-gatherers to show that it is the diversity seen among extant hunter-gatherers-both intra- and inter-population variability-that is most relevant and useful for understanding past hunter-gatherer demography.

Accurate age estimation in small-scale societies.

Precise estimation of age is essential in evolutionary anthropology, especially to infer population age structures and understand the evolution of human life history diversity. However, in small-scale societies, such as hunter-gatherer populations, time is often not referred to in calendar years, and accurate age estimation remains a challenge. We address this issue by proposing a Bayesian approach that accounts for age uncertainty inherent to fieldwork data. We developed a Gibbs sampling Markov chain Monte Carlo algorithm that produces posterior distributions of ages for each individual, based on a ranking order of individuals from youngest to oldest and age ranges for each individual. We first validate our method on 65 Agta foragers from the Philippines with known ages, and show that our method generates age estimations that are superior to previously published regression-based approaches. We then use data on 587 Agta collected during recent fieldwork to demonstrate how multiple partial age ranks coming from multiple camps of hunter-gatherers can be integrated. Finally, we exemplify how the distributions generated by our method can be used to estimate important demographic parameters in small-scale societies: here, age-specific fertility patterns. Our flexible Bayesian approach will be especially useful to improve cross-cultural life history datasets for small-scale societies for which reliable age records are difficult to acquire.

Reproductive trade-offs in extant hunter-gatherers suggest adaptive mechanism for the Neolithic expansion.

The Neolithic demographic transition remains a paradox, because it is associated with both higher rates of population growth and increased morbidity and mortality rates. Here we reconcile the conflicting evidence by proposing that the spread of agriculture involved a life history quality-quantity trade-off whereby mothers traded offspring survival for increased fertility, achieving greater reproductive success despite deteriorating health. We test this hypothesis by investigating fertility, mortality, health, and overall reproductive success in Agta hunter-gatherers whose camps exhibit variable levels of sedentarization, mobility, and involvement in agricultural activities. We conducted blood composition tests in 345 Agta and found that viral and helminthic infections as well as child mortality rates were significantly increased with sedentarization. Nonetheless, both age-controlled fertility and overall reproductive success were positively affected by sedentarization and participation in cultivation. Thus, we provide the first empirical evidence, to our knowledge, of an adaptive mechanism in foragers that reconciles the decline in health and child survival with the observed demographic expansion during the Neolithic.

Hunter-Gatherer children's close-proximity networks: Similarities and differences with cooperative and communal breeding systems.

Among vertebrates, allomothering (non-maternal care) is classified as cooperative breeding (help from sexually mature non-breeders, usually close relatives) or communal breeding (shared care between multiple breeders who are not necessarily related). Humans have been described with both labels, most frequently as cooperative breeders. However, few studies have quantified the relative contributions of allomothers according to whether they are (a) sexually mature and reproductively active and (b) related or unrelated. We constructed close-proximity networks of Agta and BaYaka hunter-gatherers. We used portable remote-sensing devices to quantify the proportion of time children under the age of 4 spent in close proximity to different categories of potential allomother. Both related and unrelated, and reproductively active and inactive, campmates had substantial involvement in children's close-proximity networks. Unrelated campmates, siblings and subadults were the most involved in both populations, whereas the involvement of fathers and grandmothers was the most variable between the two populations. Finally, the involvement of sexually mature, reproductively inactive adults was low. Where possible, we compared our findings with studies of other hunter-gatherer societies, and observed numerous consistent trends. Based on our results we discuss why hunter-gatherer allomothering cannot be fully characterised as cooperative or communal breeding.

Sedentarization and maternal childcare networks: role of risk, gender and demography.

Women cooperate over multiple domains and while research from western contexts portrays women's networks as limited in size and breadth, women receive help, particularly with childcare, from a diverse range of individuals (allomothers). Nonetheless, little exploration has occurred into why we see such diversity. Wide maternal childcare networks may be a consequence of a lack of resource accumulation in mobile hunter-gatherers-where instead households rely on risk-pooling in informal insurance networks. By contrast, when households settle and accumulate resources, they are able to retain risk by absorbing losses. Thus, the size and composition of mothers' childcare networks may depend on risk-buffering, as captured by mobile and settled households in the Agta, a Philippine foraging population with diverse lifestyles. Across 78 children, we find that childcare from grandmothers and sisters was higher in settled camps, while childcare from male kin was lower, offering little support for risk-buffering. Nonetheless, girls' workloads were increased in settled camps while grandmothers had fewer dependent children, increasing their availability. These results point to gender-specific changes associated with shifting demographics as camps become larger and more settled. Evidently, women's social networks, rather than being constrained by biology, are responsive to the changing socioecological context. This article is part of the theme issue 'Cooperation among women: evolutionary and cross-cultural perspectives'.

Wealth, health and inequality in Agta foragers.

Background and objectives: There is significant evidence from large-scale, industrial and post-industrial societies that greater income and wealth inequality is negatively associated with both population health and increasing health inequalities. However, whether such relationships are inevitable and should be expected to impact the health of small-scale societies as they become more market-integrated is less clear. Methodology: Here, using mixed-effect models, we explore the relationship between health, wealth, wealth inequality and health inequalities in a small-scale foraging population from the Philippines, the Agta. Results: Across 11 camps, we find small to moderate degrees of wealth inequality (maximal Gini Coefficient 0.44) which is highest in the most permanent camps, where individuals engage more heavily in the formal market. However, in both adults (n = 161) and children (n = 215), we find little evidence that either wealth or wealth inequality associates with ill health, except for one measure of nutritional condition-red blood cell count. Conclusions and implications: We interpret these results in the light of high levels of cooperation among the Agta which may buffer against the detrimental effects of wealth inequality documented in industrial and post-industrial societies. We observe little intergenerational wealth transmission, highlighting the fluid nature of wealth, and thus wealth inequality, particularly in mobile communities. The deterioration of nutritional status, as indicated by red blood cell counts, requires further investigation before concluding the Agta's extensive cooperation networks may be beginning to breakdown in the face of increasing inequality.

Cooperation and partner choice among Agta hunter-gatherer children: An evolutionary developmental perspective.

Examining development is essential for a full understanding of behaviour, including how individuals acquire traits and how adaptive evolutionary forces shape these processes. The present study explores the development of cooperative behaviour among the Agta, a Filipino hunter-gatherer population. A simple resource allocation game assessing both levels of cooperation (how much children shared) and patterns of partner choice (who they shared with) was played with 179 children between the ages of 3 and 18. Children were given five resources (candies) and for each was asked whether to keep it for themselves or share with someone else, and if so, who this was. Between-camp variation in children's cooperative behaviour was substantial, and the only strong predictor of children's cooperation was the average level of cooperation among adults in camp; that is, children were more cooperative in camps where adults were more cooperative. Neither age, sex, relatedness or parental levels of cooperation were strongly associated with the amount children shared. Children preferentially shared with close kin (especially siblings), although older children increasingly shared with less-related individuals. Findings are discussed in terms of their implications for understanding cross-cultural patterns of children's cooperation, and broader links with human cooperative childcare and life history evolution.

Who cares for women with children? Crossing the bridge between disciplines.

It has long been acknowledged that women with children require social support to promote their health and wellbeing, as well as that of their children. However, the dominant conceptualizations of support have been heavily influenced by Western family norms. The consequence, at best, has been to stifle our understanding of the nature and consequences of support for mothers and children. At worst, it has led to systematic discrimination negatively impacting maternal-child health. To fully engage with the complexities of social support, we must take multidisciplinary or interdisciplinary approaches spanning diverse cultural and geographical perspectives. However, multidisciplinary knowledge-processing can be challenging, and it is often unclear how different studies from different disciplines relate. To address this, we outline two epistemological frameworks-the scientific approach and Tinbergen's four questions-that can be useful tools in connecting research across disciplines. In this theme issue on 'Multidisciplinary perspectives on social support and maternal-child health', we attempt to foster multidisciplinary thinking by presenting work from a diverse range of disciplines, populations and cultures. Our hope is that these tools, along with papers in this issue, help to build a holistic understanding of social support and its consequences for mothers and their children. Overall, a multidisciplinary perspective points to how the responsibility of childrearing should not fall solely onto mothers. Indeed, this multidisciplinary issue demonstrates that successful childrearing is consistently an activity shared beyond the mother and the nuclear family: an insight that is crucial to harnessing the potential of social support to improve maternal-child health. This article is part of the theme issue 'Multidisciplinary perspectives on social support and maternal-child health'.

Relatedness within and between Agta residential groups.

Theoretical models relating to the evolution of human behaviour usually make assumptions about the kinship structure of social groups. Since humans were hunter-gatherers for most of our evolutionary history, data on the composition of contemporary hunter-gatherer groups has long been used to inform these models. Although several papers have taken a broad view of hunter-gatherer social organisation, it is also useful to explore data from single populations in more depth. Here, we describe patterns of relatedness among the Palanan Agta, hunter-gatherers from the northern Philippines. Across 271 adults, mean relatedness to adults across the population is r = 0.01 and to adult campmates is r = 0.074, estimates that are similar to those seen in other hunter-gatherers. We also report the distribution of kin across camps, relatedness and age differences between spouses, and the degree of shared reproductive interest between camp mates, a measure that incorporates affinal kinship. For both this this measure (s) and standard relatedness (r), we see no major age or sex differences in the relatedness of adults to their campmates, conditions that may reduce the potential for conflicts of interest within social groups.

Children are important too: juvenile playgroups and maternal childcare in a foraging population, the Agta.

Non-maternal carers (allomothers) are hypothesized to lighten the mother's workload, allowing for the specialized human life history including relatively short interbirth intervals and multiple dependent offspring. Here, using in-depth observational data on childcare provided to 78 Agta children (a foraging population in the northern Philippines; aged 0-6 years), we explore whether allomaternal childcare substitutes and decreases maternal childcare. We found that allomother caregiving was associated with reduced maternal childcare, but the substitutive effect varied depending on the source and type of care. Children-only playgroups consistently predicted a decrease in maternal childcare. While grandmothers were rarely available, their presence was negatively associated with maternal presence and childcare, and grandmothers performed similar childcare activities to mothers. These results underscore the importance of allomothering in reducing maternal childcare in the Agta. Our findings suggest that flexibility in childcare sources, including children-only playgroups, may have been the key to human life-history evolution. Overall, our results reinforce the necessity of a broad conceptualization of social support in human childcare. This article is part of the theme issue 'Multidisciplinary perspectives on social support and maternal-child health'.

HLA-G genetic diversity and evolutive aspects in worldwide populations.

HLA-G is a promiscuous immune checkpoint molecule. The HLA-G gene presents substantial nucleotide variability in its regulatory regions. However, it encodes a limited number of proteins compared to classical HLA class I genes. We characterized the HLA-G genetic variability in 4640 individuals from 88 different population samples across the globe by using a state-of-the-art method to characterize polymorphisms and haplotypes from high-coverage next-generation sequencing data. We also provide insights regarding the HLA-G genetic diversity and a resource for future studies evaluating HLA-G polymorphisms in different populations and association studies. Despite the great haplotype variability, we demonstrated that: (1) most of the HLA-G polymorphisms are in introns and regulatory sequences, and these are the sites with evidence of balancing selection, (2) linkage disequilibrium is high throughout the gene, extending up to HLA-A, (3) there are few proteins frequently observed in worldwide populations, with lack of variation in residues associated with major HLA-G biological properties (dimer formation, interaction with leukocyte receptors). These observations corroborate the role of HLA-G as an immune checkpoint molecule rather than as an antigen-presenting molecule. Understanding HLA-G variability across populations is relevant for disease association and functional studies.

Typologies of postnatal support and breastfeeding at two months in the UK: Response to comments by Harpur & Haddon.

We welcome the comments by Harpur and Haddon (2020) on our paper on the typologies of social support and its associations with breastfeeding at two months in a UK sample. We share their concerns around the under-acknowledged costs of breastfeeding, and the need for a truly family-centred approach to breastfeeding support. However, they are mistaken to assume we do not view breastfeeding problems as an important cost of breastfeeding: We explicitly comment on breastfeeding challenges within our paper, and such challenges are theoretically incorporated into the "costs of breastfeeding" following an evolutionary anthropological framework. They are also incorrect in their statement that we recommend breastfeeding promotion messages to wider family members. In fact, we outline in our manuscript that breastfeeding promotional messages can have negative consequences for mothers, and are unlikely to be supportive. What we do suggest is a family-centred approach that recognises that women interact with, and may be supported by, a wide range of individuals (including fathers and grandmothers); and the importance of taking a nuanced approach to support without assuming that all types of support necessarily leads to "more breastfeeding." We hope our response is useful in clarifying the key points of our paper.

Typologies of postnatal support and breastfeeding at two months in the UK.

There is extensive evidence to suggest that social support improves breastfeeding outcomes. Building on this evidence-base, public health services and interventions aiming to improve breastfeeding rates have primarily targeted informational and emotional support to mothers, reflecting an individual behaviour-change approach. However, mothers exist within a wider social network, and the characteristics of their broader support networks may be an important predictor of breastfeeding outcomes. Here we explore the typologies of postnatal support for mothers in the UK; a population with one of the lowest breastfeeding rates in Europe. Using retrospective data from an online survey (data collection period December 2017 - February 2018), we carry out a latent class regression (n = 432) to identify "clusters" of postnatal support in our data. Mothers in our sample were most likely to report receiving practical and emotional support from partners and maternal grandmothers, and breastfeeding information from health professionals. We identify three distinct typologies of postnatal support: 1) Extensive support, where mothers received support from a wide range of supporters including partners, maternal grandmothers, friends and health professionals, but mothers were the only ones to feed the infant; 2) Family support, where mothers received support from partners and maternal grandmothers, including with infant feeding, but less likely to receive support from health professionals; and 3) Low support, where mothers primarily received support from partners. 94% of women with extensive support were predicted to be breastfeeding at two months, followed by 48% of mothers in the low support group, and 13% in the family support group. Our findings highlight the complexities of family support and its potential impact on breastfeeding, as well as the significance of professional support. Overall, our results hint at the potential value for health professionals to engage with wider family in order to achieve extensive support for mothers.

Subjective Environmental Experiences and Women's Breastfeeding Journeys: A Survival Analysis Using an Online Survey of UK Mothers.

Local physical and social environmental factors are important drivers of human health and behaviour. Environmental perception has been linked with both reproduction and parenting, but links between subjective environmental experiences and breastfeeding remain unclear. Using retrospective data from an online survey of UK mothers of children aged 0-24 months, Cox-Aalen survival models test whether negative subjective environmental experiences negatively correlated with any and exclusive breastfeeding (max n = 473). Matching predictions, hazards of stopping any breastfeeding were increased, albeit non-significantly, across the five environmental measures (HR: 1.05-1.26) Hazards for stopping exclusive breastfeeding were however (non-significantly) reduced (HR: 0.65-0.87). Score processes found no significant time-varying effects. However, estimated cumulative coefficient graphs showed that the first few weeks postpartum were most susceptible to environmental influences and that contrary to our predictions, mothers with worse subjective environmental experiences were less likely to stop breastfeeding at this time. In addition, the hazard of stopping exclusive breastfeeding declined over time for mothers who thought that littering was a problem. The predicted increased hazards of stopping breastfeeding were only evident in the later stages of any breastfeeding and only for mothers who reported littering as a problem or that people tended not to know each other. Perceived harsher physical and social environmental conditions are assumed to deter women from breastfeeding, but this may not always be the case. Women's hazards of stopping breastfeeding change over time and there may be particular timepoints in their breastfeeding journeys where subjective environmental experiences play a role.

Hunter-gatherer multilevel sociality accelerates cumulative cultural evolution.

Although multilevel sociality is a universal feature of human social organization, its functional relevance remains unclear. Here, we investigated the effect of multilevel sociality on cumulative cultural evolution by using wireless sensing technology to map inter- and intraband social networks among Agta hunter-gatherers. By simulating the accumulation of cultural innovations over the real Agta multicamp networks, we demonstrate that multilevel sociality accelerates cultural differentiation and cumulative cultural evolution. Our results suggest that hunter-gatherer social structures [based on (i) clustering of families within camps and camps within regions, (ii) cultural transmission within kinship networks, and (iii) high intercamp mobility] may have allowed past and present hunter-gatherers to maintain cumulative cultural adaptation despite low population density, a feature that may have been critical in facilitating the global expansion of Homo sapiens.

Why so many Agta boys? Explaining 'extreme' sex ratios in Philippine foragers.

Male-biased sex ratios have been observed in multiple small-scale societies. Although intentional and systematic female-biased mortality has been posited as an explanation, there is often a lack of ethnographic evidence of systematic female neglect and/or infanticide. The Agta, a foraging population from the Philippines, have a skewed sex ratio of 1.29 (129 males per 100 females) aged 15 years or under. We hypothesised that this skew was not caused by greater female deaths, but due to an adaptive response, where more males were produced at birth in reaction to high male-biased extrinsic mortality. To test this hypothesis we utilise census, childcare and mortality data from 915 Agta. The Agta's sex ratio is significantly male-biased in the <1 (n = 48, 2:1) and 1-5 year (n = 170, 1.39:1) age cohorts; however, we find no evidence of systematic female neglect in patterns of childcare. Furthermore, the sex ratio decreases over cohorts, becoming balanced by the end of the juvenile period, owing to significantly higher male mortality. Taken together, these results are not supportive of female infanticide or neglect, and instead suggest an adaptive mechanism, acting in utero as a response to male-biased juvenile mortality, following Fisherian principles of equalising parental investment.