Spread of cattle led to the loss of matrilineal descent in Africa: a coevolutionary analysis

Matrilineal descent is rare in human societies that keep large livestock. However, this negative correlation does not provide reliable evidence that livestock and descent rules are functionally related, because human cultures are not statistically independent owing to their historical relationships (Galton''s problem). We tested the hypothesis that when matrilineal cultures acquire cattle they become patrilineal using a sample of 68 Bantu- and Bantoid-speaking populations from sub-Saharan Africa. We used a phylogenetic comparative method to control for Galton''s problem, and a maximum-parsimony Bantu language tree as a model of population history. We tested for coevolution between cattle and descent. We also tested the direction of cultural evolution--were cattle acquired before matriliny was lost? The results support the hypothesis that acquiring cattle led formerly matrilineal Bantu-speaking cultures to change to patrilineal or mixed descent. We discuss possible reasons for matriliny''s association with horticulture and its rarity in pastoralist societies. We outline the daughter-biased parental investment hypothesis for matriliny, which is supported by data on sex, wealth and reproductive success from two African societies, the matrilineal Chewa in Malawi and the patrilineal Gabbra in Kenya.     

Evolutionary Contributions to Solving the ��Matrilineal Puzzle��

Matriliny has long been debated by anthropologists positing either its primitive or its puzzling nature. More recently, evolutionary anthropologists have attempted to recast matriliny as an adaptive solution to modern social and ecological environments, tying together much of what was known to be associated with matriliny. This paper briefly reviews the major anthropological currents in studies of matriliny and discusses the contribution of evolutionary anthropology to this body of literature. It discusses the utility of an evolutionary framework in the context of the first independent test of Holden et al.'s 2003 model of matriliny as daughter-biased investment. It finds that historical daughter-biased transmission of land among the Mosuo is consistent with the model, whereas current income transmission is not. In both cases, resources had equivalent impacts on male and female reproduction, a result which predicts daughter-biased resource transmission given any nonzero level of paternity uncertainty. However, whereas land was transmitted traditionally to daughters, income today is invested in both sexes. Possible reasons for this discrepancy are discussed.     

Identification of visual paternity cues in humans

Understanding how individuals identify their relatives has implications for the evolution of social behaviour. Kinship cues might be based on familiarity, but in the face of paternity uncertainty and costly paternal investment, other mechanisms such as phenotypic matching may have evolved. In humans, paternal recognition of offspring and subsequent discriminative paternal investment have been linked to father–offspring facial phenotypic similarities. However, the extent to which paternity detection is impaired by environmentally induced facial information is unclear. We used 27 portraits of fathers and their adult sons to quantify the level of paternity detection according to experimental treatments that manipulate the location, type and quantity of visible facial information. We found that (i) the lower part of the face, that changes most with development, does not contain paternity cues, (ii) paternity can be detected even if relational information within the face is disrupted and (iii) the signal depends on the presence of specific information rather than their number. Taken together, the results support the view that environmental effects have little influence on the detection of paternity using facial similarities. This suggests that the cognitive dispositions enabling the facial detection of kinship relationships ignore genetic irrelevant facial information.     

Do grandparents favor granddaughters? Biased grandparental investment in UK

Differential grandparental investment in grandchildren is often explained with paternity uncertainty. The asymmetric inheritance of the sex chromosomes, especially of the X chromosomes, may also bias grandparental investment. Recent studies show that ignoring the sex of the grandchild can mask important differences in the investment patterns of the same grandparent category, but this has not been tested in contemporary societies with nationally representative data. With 17 variables from the Involved Grandparenting and Child Well-Being 2007 survey, we tested differential grandparental investment as reported by British and Welsh adolescents and compared predictions based on X-chromosomal relatedness with predictions based on paternity uncertainty. The theories are expected to differ with regard to grandmaternal investment in grandsons and granddaughters. We test whether paternal grandmothers invest (H1) more in granddaughters than in grandsons, (H2) more in granddaughters than maternal grandmothers do and (H3) less in grandsons than maternal grandmothers do. In addition, following the suggestion that paternal grandmothers may reduce sibling competition between girls and boys by harming grandsons, we study whether (H4) paternal grandmothers channel more noninvestment into grandsons than into granddaughters. The results show no convincing support for the type of sex discrimination of grandchildren that is predicted by X-chromosomal relatedness theories, but do provide support for the paternity uncertainty theory. X-chromosomal relatedness does not appear to shape grandparental behavior in developed societies.     

The coevolution of human fertility and wealth inheritance strategies.

Life history theory concerns the scheduling of births and the level of parental investment in each offspring. In most human societies the inheritance of wealth is an important part of parental investment. Patterns of wealth inheritance and other reproductive decisions, such as family size, would be expected to influence each other. Here I present an adaptive model of human reproductive decision-making, using a state-dependent dynamic model. Two decisions made by parents are considered: when to have another baby, and thus the pattern of reproduction through life; and how to allocate resources between children at the end of the parents'' life. Optimal decision rules are those that maximize the number of grandchildren. Decisions are assumed to depend on the state of the parent, which is described at any time by two variables: number of living sons, and wealth. The dynamics of the model are based on a traditional African pastoralist system, but it is general enough to approximate to any means of subsistence where an increase in the amount of wealth owned increases the capacity for future production of resources. The model is used to show that, in the unpredictable environment of a traditional pastoralist society, high fertility and a biasing of wealth inheritance to a small number of children are frequently optimal. Most such societies are now undergoing a transition to lower fertility, known as the demographic transition. The effects on fertility and wealth inheritance strategies of reducing mortality risks, reducing the unpredictability of the environment and increasing the costs of raising children are explored. Reducing mortality has little effect on completed family sizes of living children or on the wealth they inherit. Increasing the costs of raising children decreases optimal fertility and increases the inheritance left to each child at each level of wealth, and has the potential to reduce fertility to very low levels. The results offer an explanation for why wealthy families are frequently also those with the smallest number of children in heterogeneous, post-transition societies.     

Gender and Risk in a Matrifocal Caribbean Community: A View from Behavioral Ecology

Matrifocality is a feature of Caribbean communities in which mothers and adult daughters often form the household core. I argue that daughter-biased parental care underlies matrifocality. Parental investment (PI) theory predicts sex-biased care, but factors promoting daughter preference are not always clearly specified. If sons are more likely than daughters to experience unpredictable hazards, then parents may bias their efforts toward daughters. In this study, I examine gender differences in rural Dominica and test PI predictions. Men were more likely to be poor and develop alcoholism and less likely to migrate or attend high school than women were. Educational outcomes showed a Trivers-Willard effect: Boys from unfavorable family environments were less likely to receive secondary education than were other boys, but there was no association for girls. PI variables generally accounted for less variance in men's outcomes than women's, suggesting that unpredictable hazards for sons may promote daughter preference and matrifocality.     

Brothers and sisters

Data from the Kipsigis of Kenya are used to test two models for how parents invest in offspring, the Trivers-Willard and local resource competition/enhancement hypotheses. Investment is measured as age-specific survival, educational success, marital arrangements, and some components of property inheritance, permitting an evaluation of how biases persist or alter over the period of dependence. Changes through time in such biases are also examined. Despite stronger effects of wealth on the reproductive success of men than women, the survival of sons and daughters is not related to parental wealth. However, a Trivers-Willard effect characterizes educational investment: poor families show a greater concern for daughters’ (vis-à-vis sons’) schooling than do rich families, a trend that has increased over time. In regard to models of local resource competition and enhancement, men’s reproductive success decreases with number of brothers and increases with number of sisters; this pattern of competition with same-sex sibs and cooperation with opposite-sex sibs is not found among women. As predicted from these observations, parents show reduced investment in sons with a large number of brothers, and increased investment in sons with a large number of sisters. By contrast, investment in daughters is entirely unaffected by number of sisters and is influenced only in subtle ways by number of brothers. Levels of investment in relation to sibship size (irrespective of siblings’ sex) are highest for younger children of large sib sets. Discussion of the results in relation to those from other studies leads to three conclusions. First, predictive models for how investment biases vary across societies must consider a broad range of socioecological factors constraining parental options and payoffs. Second, the timing of investment biases within societies will be affected by the value of children and the costs of parental investment. Third, measures of investment appropriate for between-sex and between-class comparisons need careful attention. Each of these issues is brought to bear on the question of why, in contrast to so many other parts of the world, sex preferences are so muted in Africa.     

The evolution of matrilineal kinship organization

Matrilineal kinship organization is a human social system that emphasizes interactions between matrilineal kin, i.e. individuals related only through females. The ‘matrilineal puzzle’ refers to the potential for tension characteristic of this social system, owing to the conflict between the interests and responsibilities of men in their roles as brother/uncle versus husband/father. From an evolutionary perspective, matrilineal kinship organization is puzzling when it diverts investment of resources from the individuals who provide the potentially highest reproductive returns. I use a game-theoretic framework to investigate a particular form of matrilineal inheritance—the transfer of property from a maternal uncle to a sororal nephew. The analysis reveals two mechanisms that may make this strategy a stable evolutionary outcome. First, a polygynous male has multiple wives, and hence multiple brothers-in-law; with matrilineal inheritance, each additional brother-in-law may transfer resources to the male''s wife''s offspring, thus potentially contributing to the male''s inclusive fitness. Second, the husband of a polyandrous female is effectively ‘sharing’ paternity with other men; depending on the number of husbands, he may be better off investing in his sister''s offspring. I conclude by discussing how these results address the challenges posed by the occurrence of matrilineal kinship organization.     

Avoidance of close consanguineous inbreeding in captive groups of rhesus macaques

In three captive groups of rhesus macaques (Macaca mulatto) the intensity of inbreeding avoidance was directly correlated with the closeness of kinship. The incidence of inbreeding between matrilineal relatives was lower than expected were mating to occur randomly with regard to matrilineal affiliation. The avoidance of mating between sons and their mothers and between matrilineal sibs contributed about squally to this outcome. The incidence of mating between more remote matrilineal relatives was lower than, but could not be shown to differ significantly from, that expected at the 0.05 level of probability. Familiarity fostered by interactions among matriline members probably provides a focus for avoidance of matrilineal inbreeding, and might influence male dispersal. Results of this study are consistent with reports of female rhesus macaques' avoidance of sexual activity with matrilineal male relatives as predicted by parental investment theory The observed incidence of inbreeding between patrilineal relatives was not different from that expected, but significantly fewer of the patrilineally inbred matings involved father/daughter matings than expected were mating random. Recognition of phenotypic similarities might provide a focus for avoidance of inbreeding between fathers and their daughters, but is loss reliable than strategies for avoiding matrilineal inbreeding. Adaptations for avoiding matrilineal inbreeding that are more effective than those for avoiding patrilineal inbreeding might have evolved because the risk of patrilineal inbreeding, and hence the resulting loss in fitness, is marginal compared to that for matrilineal inbreeding in free-ranging groups. © 1995 Wiley-Liss, Inc.     

Extra-pair young in house wren broods are more likely to be male than female

Sex-allocation theory predicts that females should preferentially produce offspring of the sex with greater fitness potential. In socially monogamous animal species, extra-pair mating often increases the variance in fitness of sons relative to daughters. Thus, in situations where offspring sired by a female''s extra-pair mate(s) will typically have greater fitness potential than offspring sired by the within-pair mate, sex-allocation theory predicts that females will bias the sex of offspring sired by extra-pair mates towards male. We examined the relationship between offspring sex and paternity over six breeding seasons in an Illinois population of the house wren (Troglodytes aedon), a cavity-nesting songbird. Out of the 2345 nestlings that had both sex and paternity assigned, 350 (15%) were sired by extra-pair males. The sex ratio of extra-pair offspring, 0.534, was significantly greater than the sex ratio of within-pair offspring, 0.492, representing an increase of 8.5 per cent in the proportion of sons produced. To our knowledge, this is the first confirmed report of female birds increasing their production of sons in association with extra-pair fertilization. Our results are consistent with the oft-mentioned hypothesis that females engage in extra-pair mating to increase offspring quality.     

Altruism towards cousins

Recent research on kin investment shows a matrilateral bias as a function of paternity uncertainty. Kin investment, however, is a special case of kin altruism. We thus hypothesize that psychological adaptations have evolved to regulate cousin-directed altruism according to predictably variable levels of paternity uncertainty in different categories of cousins. We develop a formal mathematical model that predicts that individuals should be most willing to act altruistically towards their mother's sister's (MoSis) children and least willing to act altruistically towards their father's brother's (FaBro) children. Altruism towards father's sister's (FaSis) and mother's brother's (MoBro) children are predicted to fall in between. An empirical study (N=195), assessing expressed altruistic proclivities, confirmed the predictions from the model. Participants expressed willingness-to-help following the descending order: (i) MoSis children, (ii) MoBro children, (iii) FaSis children, and (iv) FaBro children. The psychological variables of emotional closeness, empathic concern and contact frequency showed precisely the same pattern across distinct cousins, providing convergent confirmation of the model. The results support the hypothesis of cousin-specific adaptations sensitive to varying probabilities of paternity uncertainty.     

Matrilateral biases in the investment of aunts and uncles

Gaulin, McBurney, and Brakeman-Wartell (1997) found that college students reported both matrilateral and sex biases in the investment of aunts and uncles (aunts invested more than uncles). They interpreted the matrilateral bias as a consequence of paternity uncertainty. We replicated that study with Orthodox Jewish college students, selected because they come from a population we presume to have higher paternity certainty than the general population. The Orthodox sample also showed matrilateral and sex biases. Comparing the two data sets, the Orthodox sample reported more investment, and slightly less matrilateral and sex biases, but the differences were not statistically significant. We did find an interaction between sex of relative and group membership, resulting from greater investment by Orthodox uncles. We interpret the results as reflecting the operation of a facultative investment mechanism whose upper limit is tuned to the maximum levels of paternity certainty found in ancestral environments. Lack of a difference in matrilateral bias between groups may result from levels of paternity certainty near to, or above, that maximum in both groups.     

Genetic relatedness to sisters' children has been underestimated

Males of many species help in the care and provisioning of offspring, and these investments often correlate with genetic relatedness. For example, many human males invest in the children of sisters, and this is especially so where men are less likely to share genes with children of wives. Although this makes qualitative sense, it has been difficult to support quantitatively. The prevailing model predicts investment in children of sisters only when paternity confidence falls below 0.268. This value is often seen as too low to be credible; so investment in sisters'' children represents an unsolved problem. I show here that the prevailing model rests on a series of restrictive assumptions that underestimate relatedness to sisters'' children. For this reason, it understates the fitness payoff to men who invest in these children. This effect can be substantial, especially in societies with low confidence in paternity. But this effect cannot be estimated solely from confidence in paternity. One must also estimate the probability that two siblings share the same father.     

Winter resource wealth drives delayed dispersal and family-group living in western bluebirds

Delayed dispersal, where offspring remain with parents beyond the usual period of dependence, is the typical route leading to formation of kin-based cooperative societies. The prevailing explanations for why offspring stay home are variation in resource wealth, in which offspring of wealthy parents benefit disproportionately by staying home, and nepotism, where the tendency for parents to be less aggressive and share food with offspring makes home a superior place to wait to breed. These hypotheses are not strict alternatives, as only wealthy parents have sufficient resources to share. In western bluebirds, Sialia mexicana, sons usually delay dispersal until after winter, gaining feeding advantages through maternal nepotism in a familial winter group. Experimentally reducing resource wealth (mistletoe) by half on winter territories caused sons to disperse in summer, even though their parents remained on the territory during the winter. Only 8% of sons remained with their parents on mistletoe-removal territories compared to 50% of sons on control territories (t(9,10)=3.33, p<0.005). This study is the first to demonstrate that experimentally reducing wealth of a natural food resource reduces delayed dispersal, facilitating nepotism and family-group living. The results clarify the roles of year-round residency, resource limitation and relative wealth outside the breeding season in facilitating the formation of kin-based cooperative societies.     

Male genital mutilation: an adaptation to sexual conflict

Male genital mutilation (MGM) takes several forms and occurs in about 25% of societies. This behavior has puzzled anthropologists, doctors and theologians for centuries, and presents an evolutionary challenge since it involves dangerous and costly surgery. I suggest that MGM is likely to reduce insemination efficiency, reducing a man's capacity for extra-pair fertilizations by impairing sperm competition. MGM may therefore represent a hard-to-fake signal of a man's reduced ability to challenge the paternity of older men who are already married. Men who display this signal of sexual obedience may gain social benefits if married men are selected to offer social trust and investment preferentially to peers who are less threatening to their paternity. Clitoridectomy and vaginal infibulation serve a parallel signaling function in women, increasing a husband's paternity certainty and garnering his increased investment. Especially in societies where paternity uncertainty and reproductive conflict are high, the social benefits of MGM as a signal may outweigh its costs. This ‘sexual conflict’ hypothesis predicts that MGM should be associated with polygyny, particularly when co-wives reside far apart, and that MGM should reduce the frequency of extramarital sex. MGM rituals should facilitate access to social benefits; they should be highly public, watched mainly by men, and performed by a nonrelative. I found support for these six predictions in two cross-cultural samples. I also examined an alternative hypothesis suggesting that MGM signals group commitment for collective action, particularly inter-societal warfare. Although other forms of male scarification fit this model, the distribution of MGM is not predicted by frequency of inter-societal warfare.     

Confidence of paternity,divorce, and investment in children by Albuquerque men

Using a sample of men living in Albuquerque, NM, we examined the relationship between paternity confidence and men's investment in children. In humans, men may reduce their investment in a child in two ways: indirectly, by ending their relationship with the child's mother and ceasing to cohabit with the child (e.g., divorce), and directly, by allocating less time and fewer resources to the child. In this article, we tested two hypotheses regarding the effect of paternity confidence on investment in children: (1) men will be more likely to divorce women if they suspect or are sure that they are not the father of their wife's child, and (2) controlling for divorce, men will reduce direct investments in low paternity confidence children relative to high paternity confidence children. The first hypothesis was supported by the data. The second hypothesis was supported for two out of three measures of paternal investment we examined; low paternity confidence reduces the time men spend with a child in a group with other children or adults, and it reduces extensive involvement with the child's educational progress; there was no effect of paternity confidence on the amount of time men spend with children in one-on-one interactions. We also examined the effects of unstated paternity confidence (e.g., when men decline to answer the question) on divorce and paternal investment. Overall, the results suggested that paternity confidence plays an important role in shaping men's relationships with women and with their putative genetic children.     

Offspring sex ratios in tree swallows: females in better condition produce more sons

Organisms are expected to adjust the sex ratio of their offspring in relation to the relative fitness benefits of sons and daughters. We used a molecular sexing technique that amplifies an intron of the CHD1 gene in birds to examine the sex ratio at egg-laying in socially monogamous tree swallows (Tachycineta bicolor). We examined all individuals in 40 broods (210 young), including all unhatched eggs and nestlings. Thus, the sex ratio we measured was the same as the sex ratio at laying. Overall, the mean sex ratio per brood (+/- SD) was biased significantly towards males (57 +/- 2% male). Within broods, male-biased sex ratios were associated with females in better body condition, and these females were more likely to produce sons in better condition. Tree swallows have one of the highest known levels of extra-pair paternity in birds (38-76% extra-pair young), and, as a consequence, variance in male reproductive success is greater than that of females. Thus, in tree swallows, investment in sons has the potential for higher fitness returns than investment in daughters, assuming that sons in better condition have greater reproductive success.     

Matrilateral biases in the investment of aunts and uncles

In a study of the kin investment of aunts and uncles we show that the laterality effect expected as a result of paternity uncertainty is statistically reliable but somewhat smaller than the sex effect. Matrilateral aunts invest significantly more than patrilateral aunts, and the same is true for uncles. Regardless of laterality, however, aunts invest significantly more than uncles. Multivariate controls show that the matrilateral bias is fully independent of any age or distance confounds that might result from sex differences in age at marriage or dispersal. We discuss our results in relation to recent findings on the kin investment of grandparents (Euler and Weitzel 1996). In addition, we propose a simple method for estimating the level of paternity uncertainty from kin investment data; application of this method to our data on aunts and uncles suggests that between 13% and 20% of children are not the offspring of their putative father. Our parallel analyses of Euler and Weitzel’s (1996) data on grandparental investment suggest a similar estimate, that paternity uncertainty lies between 9% and 17%.     

In poor families, mothers' milk is richer for daughters than sons: A test of Trivers-Willard hypothesis in agropastoral settlements in Northern Kenya

The Trivers–Willard hypothesis predicts the unequal parental investment between daughters and sons, depending on maternal condition and offspring reproductive potential. Specifically, in polygynous populations where males have higher reproductive variance than females, it predicts that mothers in good condition will invest more in sons, whereas mothers in poor condition will invest more in daughters. Previous studies testing this hypothesis focused on behavioral investment, whereas few examined biological investment. This study investigates the Trivers–Willard hypothesis on both behavioral and biological parental investment by examining breastfeeding frequencies and breast milk fat concentrations. Data from exclusively breastfeeding mothers in Northern Kenya were used to test hypotheses: Economically sufficient mothers will breastfeed sons more frequently than daughters, whereas poor mothers will breastfeed daughters more frequently than sons, and economically sufficient mothers will produce breast milk with higher fat concentration for sons than daughters, whereas poor mothers will produce breast milk with higher fat concentration for daughters than sons. Linear regression models were applied, using breastfeeding frequency or log‐transformed milk fat as the dependent variable, and offspring's sex (son = 1/daughter = 0), socioeconomic status (higher = 1/lower = 0), and the sex‐wealth interaction as the predictors, controlling for covariates. Our results only supported the milk fat hypothesis: infant's sex and socioeconomic status interacted (P = 0.014, n = 72) in their relation with milk fat concentration. The model estimated that economically sufficient mothers produced richer milk for sons than daughters (2.8 vs. 0.6 gm/dl) while poor mothers produced richer milk for daughters than sons (2.6 vs. 2.3 gm/dl). Further research on milk constituents in relation to offspring's sex is warranted. Am J Phys Anthropol , 2012. © 2012 Wiley Periodicals, Inc.