Correction: Betel quid use is associated with anemia among both men and women in Matlab, Bangladesh.

[This corrects the article DOI: 10.1371/journal.pgph.0001677.].

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.

Market integration, income inequality, and kinship system among the Mosuo of China.

Increased access to defensible material wealth is hypothesised to escalate inequality. Market integration, which creates novel opportunities in cash economies, provides a means of testing this hypothesis. Using demographic data collected from 505 households among the matrilineal and patrilineal Mosuo in 2017, we test whether market integration is associated with increased material wealth, whether increased material wealth is associated with wealth inequality, and whether being in a matrilineal vs. patrilineal kinship system alters the relationship between wealth and inequality. We find evidence that market integration, measured as distance to the nearest source of tourism and primary source of household income, is associated with increased household income and 'modern' asset value. Both village-level market integration and mean asset value were associated negatively, rather than positively, with inequality, contrary to predictions. Finally, income, modern wealth and inequality were higher in matrilineal communities that were located closer to the centre of tourism and where tourism has long provided a relatively stable source of income. However, we also observed exacerbated inequality with increasing farm animal value in patriliny. We conclude that the forces affecting wealth and inequality depend on local context and that the importance of local institutions is obscured by aggregate statistics drawn from modern nation states.

Gender disparities in material and educational resources differ by kinship system.

Contemporary inequality exists at an unprecedented scale. Social scientists have emphasized the role played by material wealth in driving its escalation. Evolutionary anthropologists understand the drive to accumulate material wealth as one that is coupled ultimately to increasing reproductive success. Owing to biological caps on reproduction for women, the efficiency of this conversion can differ by gender, with implications for understanding the evolution of gender disparities in resource accumulation. Efficiency also differs according to the type of resources used to support reproductive success. In this paper, we review evolutionary explanations of gender disparities in resources and investigate empirical evidence to support or refute those explanations among matrilineal and patrilineal subpopulations of ethnic Chinese Mosuo, who share an ethnolinguistic identity, but differ strikingly in terms of institutions and norms surrounding kinship and gender. We find that gender differentially predicts income and educational attainment. Men were more likely to report income than women; amounts earned were higher for men overall, but the difference between men and women was minimal under matriliny. Men reported higher levels of educational attainment than women, unexpectedly more so in matrilineal contexts. The results reveal nuances in how biology and cultural institutions affect gender disparities in wealth. This article is part of the theme issue 'Evolutionary ecology of inequality'.

Betel quid use is associated with anemia among both men and women in Matlab, Bangladesh.

Anemia accounts for 8.8% of total disability burden worldwide. Betel quid use among pregnant women has been found to increase anemia risk. Betel quid is prepared by wrapping the betel (or areca) nut, with spices and other additions, in betel or tobacco leaf and it is chewed or placed in the mouth. We explored the association between betel quid use and anemia among men and non-pregnant women. We collected data from a random sample of women and their husbands in Matlab, Bangladesh. Participants reported their current betel quid use and individual characteristics. We assessed hemoglobin (a biomarker of anemia) with a hemoglobinometer and soluble transferrin receptor (a biomarker of iron deficiency) and C-reactive protein (a biomarker of inflammation) in dried blood spots via enzyme immunoassay. We estimated logistic regression models to evaluate the association between betel quid use and anemia and structural equation models (SEM) to evaluate mediating roles of iron deficiency and elevated inflammation. A total of 1133 participants (390 men and 743 non-pregnant women) were included. After controlling for important confounders, any betel quid use was positively associated with anemia among men (OR: 1.80; 95% CI: 1.12, 2.89). Among women, betel quid use was associated with anemia only among the most frequent users (OR: 1.62; 95% CI: 1.03, 2.53). SEM did not reveal indirect paths through inflammation or iron deficiency. Betel quid use may contribute to the burden of anemia among adults in Bangladesh. Our findings suggest the burden of disease attributed to betel quid use has been underestimated.

Does gender structure social networks across domains of cooperation? An exploration of gendered networks among matrilineal and patrilineal Mosuo.

Cooperative networks are essential features of human society. Evolutionary theory hypothesizes that networks are used differently by men and women, yet the bulk of evidence supporting this hypothesis is based on studies conducted in a limited range of contexts and on few domains of cooperation. In this paper, we compare individual-level cooperative networks from two communities in Southwest China that differ systematically in kinship norms and institutions-one matrilineal and one patrilineal-while sharing an ethnic identity. Specifically, we investigate whether network structures differ based on prevailing kinship norms and type of gendered cooperative activity, one woman-centred (preparation of community meals) and one man-centred (farm equipment lending). Our descriptive results show a mixture of 'feminine' and 'masculine' features in all four networks. The matrilineal meals network stands out in terms of high degree skew. Exponential random graph models reveal a stronger role for geographical proximity in patriliny and a limited role of affinal relatedness across all networks. Our results point to the need to consider domains of cooperative activity alongside gender and cultural context to fully understand variation in how women and men leverage social relationships toward different ends. This article is part of the theme issue 'Cooperation among women: evolutionary and cross-cultural perspectives'.

Tradeoffs in milk immunity affect infant infectious disease risk.

Background and objectives: The human immune system has evolved to balance protection against infection with control of immune-mediated damage and tolerance of commensal microbes. Such tradeoffs between protection and harm almost certainly extend to the immune system of milk. Methodology: Among breastfeeding mother-infant dyads in Kilimanjaro, Tanzania, we characterized in vitro proinflammatory milk immune responses to Salmonella enterica (an infectious agent) and Escherichia coli (a benign target) as the increase in interleukin-6 after 24 h of incubation with each bacterium. We characterized incident infectious diseases among infants through passive monitoring. We used Cox proportional hazards models to describe associations between milk immune activity and infant infectious disease. Results: Among infants, risk for respiratory infections declined with increasing milk in vitro proinflammatory response to S. enterica (hazard ratio [HR]: 0.68; 95% confidence interval [CI]: 0.54, 0.86; P: 0.001), while risk for gastrointestinal infections increased with increasing milk in vitro proinflammatory response to E. coli (HR: 1.44; 95% CI: 1.05, 1.99; P: 0.022). Milk proinflammatory responses to S. enterica and E. coli were positively correlated (Spearman's rho: 0.60; P: 0.000). Conclusions and implications: These findings demonstrate a tradeoff in milk immune activity: the benefits of appropriate proinflammatory activity come at the hazard of misdirected proinflammatory activity. This tradeoff is likely to affect infant health in complex ways, depending on prevailing infectious disease conditions. How mother-infant dyads optimize proinflammatory milk immune activity should be a central question in future ecological-evolutionary studies of the immune system of milk.

Social support, nutrition and health among women in rural Bangladesh: complex tradeoffs in allocare, kin proximity and support network size.

Malnutrition among women of reproductive age is a significant public health concern in low- and middle-income countries. Of particular concern are undernutrition from underweight and iron deficiency, along with overweight and obesity, all of which have negative health consequences for mothers and children. Accumulating evidence suggests that risk for poor nutritional outcomes may be mitigated by social support, yet how social support is measured varies tremendously and its effects likely vary by age, kinship and reproductive status. We examine the effects of different measures of social support on weight and iron nutrition among 677 randomly sampled women from rural Bangladesh. While we find that total support network size mitigates risk for underweight, other results point to a potential tradeoff in the effects of kin proximity, with nearby adult children associated with both lower risk for underweight and obesity and higher risk for iron deficiency and anaemia. Social support from kin may then enhance energy balance but not diet quality. Results also suggest that a woman's network of caregivers might reflect their greater need for help, as those who received more help with childcare and housework had worse iron nutrition. Overall, although some findings support the hypothesis that social support can be protective, others emphasize that social relationships often have neutral or negative effects, illustrating the kinds of tradeoffs expected from an evolutionary perspective. The complexities of these effects deserve attention in future work, particularly within public health, where what is defined as 'social support' is often assumed to be positive. This article is part of the theme issue 'Multidisciplinary perspectives on social support and maternal-child health'.

Matriliny reverses gender disparities in inflammation and hypertension among the Mosuo of China.

Women experience higher morbidity than men, despite living longer. This is often attributed to biological differences between the sexes; however, the majority of societies in which these disparities are observed exhibit gender norms that favor men. We tested the hypothesis that female-biased gender norms ameliorate gender disparities in health by comparing gender differences in inflammation and hypertension among the matrilineal and patrilineal Mosuo of China. Widely reported gender disparities in health were reversed among matrilineal Mosuo compared with patrilineal Mosuo, due to substantial improvements in women's health, with no concomitant detrimental effects on men. These findings offer evidence that gender norms limiting women's autonomy and biasing inheritance toward men adversely affect the health of women, increasing women's risk for chronic diseases with tremendous global health impact.

High-altitude adaptations mitigate risk for hypertension and diabetes-associated anemia.

BACKGROUND: Human populations native to high altitude exhibit numerous genetic adaptations to hypobaric hypoxia. Among Tibetan plateau peoples, these include increased vasodilation and uncoupling of erythropoiesis from hypoxia. OBJECTIVE/METHODS: We tested the hypothesis that these high-altitude adaptations reduce risk for hypertension and diabetes-associated anemia among the Mosuo, a Tibetan-descended population in the mountains of Southwest China that is experiencing rapid economic change and increased chronic disease risk. RESULTS: Hypertension was substantially less common among Mosuo than low-altitude Han populations, and models fit to the Han predicted higher probability of hypertension than models fit to the Mosuo. Diabetes was positively associated with anemia among the Han, but not the Mosuo. CONCLUSION: The Mosuo have lower risk for hypertension and diabetes-associated anemia than the Han, supporting the hypothesis that high-altitude adaptations affecting blood and circulation intersect with chronic disease processes to lower risk for these outcomes. As chronic diseases continue to grow as global health concerns, it is important to investigate how they may be affected by local genetic adaptations.

Patterns of paternal investment predict cross-cultural variation in jealous response.

Long-lasting, romantic partnerships are a universal feature of human societies, but almost as ubiquitous is the risk of instability when one partner strays. Jealous response to the threat of infidelity is well studied, but most empirical work on the topic has focused on a proposed sex difference in the type of jealousy (sexual or emotional) that men and women find most upsetting, rather than on how jealous response varies1,2. This stems in part from the predominance of studies using student samples from industrialized populations, which represent a relatively homogenous group in terms of age, life history stage and social norms3,4. To better understand variation in jealous response, we conducted a 2-part study in 11 populations (1,048 individuals). In line with previous work, we find a robust sex difference in the classic forced-choice jealousy task. However, we also show substantial variation in jealous response across populations. Using parental investment theory, we derived several predictions about what might trigger such variation. We find that greater paternal investment and lower frequency of extramarital sex are associated with more severe jealous response. Thus, partner jealousy appears to be a facultative response, reflective of the variable risks and costs of men's investment across societies.

Differences between sons and daughters in the intergenerational transmission of wealth.

Persistent interest lies in gender inequality, especially with regard to the favouring of sons over daughters. Economists are concerned with how privilege is transmitted across generations, and anthropologists have long studied sex-biased inheritance norms. There has, however, been no focused cross-cultural investigation of how parent-offspring correlations in wealth vary by offspring sex. We estimate these correlations for 38 wealth measures, including somatic and relational wealth, from 15 populations ranging from hunter-gatherers to small-scale farmers. Although small sample sizes limit our statistical power, we find no evidence of ubiquitous male bias, at least as inferred from comparing parent-son and parent-daughter correlations. Rather we find wide variation in signatures of sex bias, with evidence of both son and daughter-biased transmission. Further, we introduce a model that helps pinpoint the conditions under which simple mid-point parent-offspring wealth correlations can reveal information about sex-biased parental investment. Our findings are relevant to the study of female-biased kinship by revealing just how little normative descriptors of kinship systems, such as patrilineal inheritance, capture intergenerational correlations in wealth, and how variable parent-son and parent-daughter correlations can be. This article is part of the theme issue 'The evolution of female-biased kinship in humans and other mammals'.

The expendable male hypothesis.

Matriliny is a system of kinship in which descent and inheritance are conferred along the female line. The theoretically influential concept of the matrilineal puzzle posits that matriliny poses special problems for understanding men's roles in matrilineal societies. Ethnographic work describes the puzzle as the tension experienced by men between the desire to exert control over their natal kin (i.e. the lineage to which they belong) and over their affinal kin (i.e. their spouses and their biological children). Evolutionary work frames the paradox as one resulting from a man investing in his nieces and nephews at the expense of his own biological offspring. In both cases, the rationale for the puzzle rests on two fundamental assumptions: (i) that men are in positions of authority over women and over resources; and (ii) that men are interested in the outcomes of parenting. In this paper, we posit a novel hypothesis that suggests that certain ecological conditions render men expendable within local kinship configurations, nullifying the above assumptions. This arises when (i) women, without significant assistance from men, are capable of meeting the subsistence needs of their families; and (ii) men have little to gain from parental investment in children. We conclude that the expendable male hypothesis may explain the evolution of matriliny in numerous cases, and by noting that female-centred approaches that call into doubt assumptions inherent to male-centred models of kinship are justified in evolutionary perspective. This article is part of the theme issue 'The evolution of female-biased kinship in humans and other mammals'.

The evolution of female-biased kinship in humans and other mammals.

Female-biased kinship (FBK) arises in numerous species and in diverse human cultures, suggesting deep evolutionary roots to female-oriented social structures. The significance of FBK has been debated for centuries in human studies, where it has often been described as difficult to explain. At the same time, studies of FBK in non-human animals point to its apparent benefits for longevity, social complexity and reproduction. Are female-biased social systems evolutionarily stable and under what circumstances? What are the causes and consequences of FBK? The purpose of this theme issue is to consolidate efforts towards understanding the evolutionary significance and stability of FBK in humans and other mammals. The issue includes broad theoretical and empirical reviews as well as specific case studies addressing the social and ecological correlates of FBK across taxa, time and space. It leverages a comparative approach to test existing hypotheses and presents novel arguments that aim to expand our understanding of how males and females negotiate kinship across diverse contexts in ways that lead to the expression of female biases in kinship behaviour and social structure. This article is part of the theme issue 'The evolution of female-biased kinship in humans and other mammals'.

Adopted daughters and adopted daughters-in-law in Taiwan: a mortality analysis.

Adoption is sometimes considered paradoxical from an evolutionary perspective because the costs spent supporting an adopted child would be better spent on rearing one's own. Kin selection theory is commonly used to solve this paradox, because the adoption of closely related kin contributes to the inclusive fitness of the adoptive parent. In this paper, we perform a novel test of kin selection theory in the context of adoption by asking whether adopted daughters-in-law, who contribute directly (i.e. genealogically) to the perpetuation of their adoptive families' lineages, experience lower mortality than daughters adopted for other purposes in historical Taiwan. We show that both classes of adopted daughter suffer lower mortality than biological daughters, but that the protective effect of adoption is stronger among daughters who were not adopted with the intention of perpetuating the family lineage. We speculate as to the possible benefits of such a pattern and emphasize the need to move beyond typological definitions of adoption to understand the specific costs and benefits involved in different forms of caring for others' children.

Offspring sex preferences among patrilineal and matrilineal Mosuo in Southwest China revealed by differences in parity progression.

Son preference predominates in China, yet there are patterned exceptions to this rule. In this paper, we test whether lineality (patrilineal versus matrilineal inheritance and descent) is associated with son versus daughter preference among the ethnic Mosuo (Na) of Southwest China. Our results show (i) an increased probability of continued fertility among matrilineal women after having a son compared with a daughter and (ii) an increased probability of continued fertility among patrilineal women after having a daughter compared with a son. These results are consistent with son preference among patrilineal Mosuo and more muted daughter preference among the matrilineal Mosuo. Furthermore, we show (iii) the lowest probability of continued fertility at parity 2 once women have one daughter and one son across both systems, suggesting that preferences for at least one of each sex exist alongside preferences for the lineal sex. The Mosuo are the only known small-scale society in which two kinship systems distinguish sub-groups with many otherwise shared cultural characteristics. We discuss why this, in conjunction with differences in subsistence, may shed light on the evolutionary underpinnings of offspring sex preferences.

Modernizing Evolutionary Anthropology : Introduction to the Special Issue.

Evolutionary anthropology has traditionally focused on the study of small-scale, largely self-sufficient societies. The increasing rarity of these societies underscores the importance of such research yet also suggests the need to understand the processes by which such societies are being lost-what we call "modernization"-and the effects of these processes on human behavior and biology. In this article, we discuss recent efforts by evolutionary anthropologists to incorporate modernization into their research and the challenges and rewards that follow. Advantages include that these studies allow for explicit testing of hypotheses that explore how behavior and biology change in conjunction with changes in social, economic, and ecological factors. In addition, modernization often provides a source of "natural experiments" since it may proceed in a piecemeal fashion through a population. Challenges arise, however, in association with reduced variability in fitness proxies such as fertility, and with the increasing use of relatively novel methodologies in evolutionary anthropology, such as the analysis of secondary data. Confronting these challenges will require careful consideration but will lead to an improved understanding of humanity. We conclude that the study of modernization offers the prospect of developing a richer evolutionary anthropology, by encompassing ultimate and proximate explanations for behavior expressed across the full range of human societies.

The evolution of inequality.

Understanding how systems of political and economic inequality evolved from relatively egalitarian origins has long been a focus of anthropological inquiry. Many hypotheses have been suggested to link socio-ecological features with the rise and spread of inequality, and empirical tests of these hypotheses in prehistoric and extant societies are increasing. In this review, we synthesize several streams of theory relevant to understanding the evolutionary origins, spread, and adaptive significance of inequality. We argue that while inequality may be produced by a variety of localized processes, its evolution is fundamentally dependent on the economic defensibility and transmissibility of wealth. Furthermore, these properties of wealth could become persistent drivers of inequality only following a shift to a more stable climate in the Holocene. We conclude by noting several key areas for future empirical research, emphasizing the need for more analyses of contemporary shifts toward institutionalized inequality as well as prehistoric cases.