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.

Mothers with higher twinning propensity had lower fertility in pre-industrial Europe.

Historically, mothers producing twins gave birth, on average, more often than non-twinners. This observation has been interpreted as twinners having higher intrinsic fertility - a tendency to conceive easily irrespective of age and other factors - which has shaped both hypotheses about why twinning persists and varies across populations, and the design of medical studies on female fertility. Here we show in >20k pre-industrial European mothers that this interpretation results from an ecological fallacy: twinners had more births not due to higher intrinsic fertility, but because mothers that gave birth more accumulated more opportunities to produce twins. Controlling for variation in the exposure to the risk of twinning reveals that mothers with higher twinning propensity - a physiological predisposition to producing twins - had fewer births, and when twin mortality was high, fewer offspring reaching adulthood. Twinning rates may thus be driven by variation in its mortality costs, rather than variation in intrinsic fertility.

Effects of Individual Mortality Experience on Out-of-Wedlock Fertility in Eighteenth- and Nineteenth-Century Krummhörn, Germany.

Life history theory predicts that exposure to high mortality in early childhood leads to faster and riskier reproductive strategies. Individuals who grew up in a high mortality regime will not overly wait until they find a suitable partner and form a stable union because premature death would prevent them from reproducing. Cox proportional hazard models were used to determine whether women who experienced sibling death during early childhood (0-5 years) reproduced earlier and were at an increased risk of giving birth to an illegitimate child, with illegitimacy serving as a proxy for risky sexual behavior. Furthermore, we investigate whether giving birth out of wedlock is influenced by individual mortality experience or by more promiscuous sexual behavior that is clustered in certain families. Models are fitted on pedigree data from the eighteenth- and nineteenth-century Krummhörn population in Germany. The results show a relationship between sibling death in early childhood and the risk of reproducing out of wedlock, and reproductive timing. The risk of giving birth out of wedlock is linked to individual mortality experience rather than to family-level effects. In contrast, adjustments in connubial reproductive timing are influenced more by family-level effects than by individual mortality experience.

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'.

Greater wealth inequality, less polygyny: rethinking the polygyny threshold model.

Monogamy appears to have become the predominant human mating system with the emergence of highly unequal agricultural populations that replaced relatively egalitarian horticultural populations, challenging the conventional idea-based on the polygyny threshold model-that polygyny should be positively associated with wealth inequality. To address this polygyny paradox, we generalize the standard polygyny threshold model to a mutual mate choice model predicting the fraction of women married polygynously. We then demonstrate two conditions that are jointly sufficient to make monogamy the predominant marriage form, even in highly unequal societies. We assess if these conditions are satisfied using individual-level data from 29 human populations. Our analysis shows that with the shift to stratified agricultural economies: (i) the population frequency of relatively poor individuals increased, increasing wealth inequality, but decreasing the frequency of individuals with sufficient wealth to secure polygynous marriage, and (ii) diminishing marginal fitness returns to additional wives prevent extremely wealthy men from obtaining as many wives as their relative wealth would otherwise predict. These conditions jointly lead to a high population-level frequency of monogamy.

When the mother-in-law is just as good-Differential mortality of reproductive females by family network composition.

Motivated by the cooperative breeding hypothesis, we investigate the effect of having kin on the mortality of reproductive women based on family reconstitutions for the Krummhörn region (East Frisia, Germany, 1720-1874). We rely on a combination of Cox clustered hazard models and hazard models stratified at the family level. In order to study behavior-related effects, we run a series of models in which only kin who lived in the same parish are considered. To investigate structural, non-behavior-related effects, we run a different model series that include all living kin, regardless their spatial proximity. We find that women of reproductive age who had a living mother had a reduced mortality risk. It appears that having living sisters had an ambivalent impact on women's mortality: i.e., depending on the socioeconomic status of the family, the effect of having living sisters ranged between representing a source of competition and representing a source of support. Models which are clustered at the family level suggest that the presence of a living mother-in-law was associated with reduced mortality among her daughters-in-law especially among larger-scale farm families. We interpret this finding as a consequence of augmented consanguineous marriages among individuals of higher social strata. For instance, in first cousin marriages, the mother-in-law could also be a biological aunt. Thus, it appears that among the wealthy elite, the genetic in-law conflict was neutralized to some extent by family solidarity. This result further suggests that the tipping point of the female trade-off between staying with the natal family and leaving the natal family to join an economically well-established in-law family might have been reached very quickly among women living under the socioeconomic conditions of the Krummhörn region.

Older fathers' children have lower evolutionary fitness across four centuries and in four populations.

Higher paternal age at offspring conception increases de novo genetic mutations. Based on evolutionary genetic theory we predicted older fathers' children, all else equal, would be less likely to survive and reproduce, i.e. have lower fitness. In sibling control studies, we find support for negative paternal age effects on offspring survival and reproductive success across four large populations with an aggregate N > 1.4 million. Three populations were pre-industrial (1670-1850) Western populations and showed negative paternal age effects on infant survival and offspring reproductive success. In twentieth-century Sweden, we found minuscule paternal age effects on survival, but found negative effects on reproductive success. Effects survived tests for key competing explanations, including maternal age and parental loss, but effects varied widely over different plausible model specifications and some competing explanations such as diminishing paternal investment and epigenetic mutations could not be tested. We can use our findings to aid in predicting the effect increasingly older parents in today's society will have on their children's survival and reproductive success. To the extent that we succeeded in isolating a mutation-driven effect of paternal age, our results can be understood to show that de novo mutations reduce offspring fitness across populations and time periods.

Human agricultural economy is, and likely always was, largely based on kinship - Why?

We question the sequence of evolutionary transitions leading to ultrasociality in humans proposed by Gowdy & Krall. Evidence indicates that families are, and likely always have been, the primary productive units in human agricultural economies, suggesting that genetic relatedness is key to understanding when the suppression of individual autonomy to the benefit of subsistence groups, that is, extended families, evolved.

Conditional grandmother effects on age at marriage, age at first birth, and completed fertility of daughters and daughters-in-law in historical Krummhörn.

Based on historical data pertaining to the Krummhörn population (eighteenth and nineteenth centuries, Germany), we compared reproductive histories of mothers according to whether the maternal grandmother (MGM) or the paternal grandmother (PGM) or neither of them was resident in the parents' parish at the time of the mother's first birth. In contrast to effects of PGMs, we discovered conditional differences in the MGM's effects between landless people and wealthier, commercial farmers. Our data indicate that the presence of the MGM only lowers the woman's age at marriage (AAM) and her age at the birth of her first child (AFB) in the case of landless families. However, among commercial farmers, who can generally be characterized by a lower AAM and AFB, we found opposite tendencies for the MGM's effect leading to a relatively small delay in AAM and AFB. Moreover, we also analyzed differences in the completed fertility (i.e., children ever born: CEB). Results indicate that landless families in general do have fewer CEB compared with commercial farmers except for those families in which the MGM has been present. Emphasizing that the adaptiveness of investment decisions should depend on the interaction of genetic, lineage-specific (intrinsic) and ecologically imposed (extrinsic) constraints, we conclude that kin strategies consequently address different fitness components under different conditions.

Grandma plays favourites: X-chromosome relatedness and sex-specific childhood mortality.

Biologists use genetic relatedness between family members to explain the evolution of many behavioural and developmental traits in humans, including altruism, kin investment and longevity. Women's post-menopausal longevity in particular is linked to genetic relatedness between family members. According to the 'grandmother hypothesis', post-menopausal women can increase their genetic contribution to future generations by increasing the survivorship of their grandchildren. While some demographic studies have found evidence for this, others have found little support for it. Here, we re-model the predictions of the grandmother hypothesis by examining the genetic relatedness between grandmothers and grandchildren. We use this new model to re-evaluate the grandmother effect in seven previously studied human populations. Boys and girls differ in the per cent of genes they share with maternal versus paternal grandmothers because of differences in X-chromosome inheritance. Here, we demonstrate a relationship between X-chromosome inheritance and grandchild mortality in the presence of a grandmother. With this sex-specific and X-chromosome approach to interpreting mortality rates, we provide a new perspective on the prevailing theory for the evolution of human female longevity. This approach yields more consistent support for the grandmother hypothesis, and has implications for the study of human evolution.

Intrafamilial resource competition and mate competition shaped social-group-specific natal dispersal in the 18th and 19th century Krummhörn population.

We describe the natal dispersal patterns of the Krummhörn population as the outcome of intrafamilial competition. Depending on the affiliation with a specific social group and the sex of the individual, this competition is driven by different factors and obeys a different functional logic: The dispersal patterns of the daughters of landless workers allow a mate competition scenario to be detected, whereas the dispersal behavior of the children of farmers (especially of the sons) is driven by a resource competition scenario.

Toward an evolutionary psychology of religiosity.

How can the evolution of religiosity be explained? To answer this question, we attempt to develop an understanding of the psychological domains underlying religious behaviour. We see four evolved domains, the sum and interaction of which constitute religiosity, namely: mysticism, ethics, myths and rituals. Even if the individual content, accents and implementations differ in each specific religion, they nevertheless derive from evolved Darwinian algorithms that are species-specific adaptations of homo sapiens. Mysticism. Intuitive ontologies are the basis for mystical experiences. Usually they serve to classify reality into animate and inanimate objects, animals or plants, for example. For a variety of psychological reasons, supernatural experiences result from a mixture of different ontological categories. Ethics. The basis for ethics lies in the social competency of human beings. Ethics is founded on the concept of social exchange ("social-contract algorithm") with its ideas about reciprocity, fairness, justice, cheater detection, in-group/out-group differentiation, etc. Myths. The basis for myths is the "language instinct". We interpret myths as the verbal expression of the cognitive content of those individual modules that constitute the belief system. Above all, myths document the experience and processing of contingency and thus help social bonding. Rituals. Rituals are based on the handicap principle. By making certain symbols and acts more expensive, they signal commitment for a reliable in-group morale. In conclusion, we argue that human religiosity emerges from a cognitive interaction between these four domains. Religiosity processes contingencies and enhances co-operation through social bonding, norm setting and cheater detection. It fulfils those functions for which the mental modules of its four domains have evolved so that we feel it appears to be justified to attribute to religiosity the evolutionary status of an adaptation.