Historical Mortality Dynamics on the Baja California Peninsula : Marriage, Mining Booms, Migration, and Infectious Disease.

Historical demographic research shows that the factors influencing mortality risk are labile across time and space. This is particularly true for datasets that span societal transitions. Here, we seek to understand how marriage, migration, and the local economy influenced mortality dynamics in a rapidly changing environment characterized by high in-migration and male-biased sex ratios. Mortality records were extracted from a compendium of historical vital records for the Baja California peninsula (Mexico). Our sample consists of 1,201 mortality records spanning AD 1835-1900. Findings from Cox proportional hazard models indicate that (1) marriage was associated with a protective effect for both sexes; (2) residing in a mining town was associated with higher mortality for men, but not women; (3) migration was associated with decreased mortality risk for women, but not men; and (4) the risk of mortality increased in the face of infectious disease, but decreased over time. Despite the early initiation of reproduction for women, marriage had a protective effect, likely because marriage linked women to resources. Although mining boomtowns were associated with elevated risk factors generally, only men experienced greater mortality risk, likely due to dangerous working conditions that women did not experience. Last, female, but not male, migrants experienced greater longevity, possibly because exposure to harsh labor conditions eroded the protective effect of selection bias for men. Together, these results shed light on an understudied historical population and broaden our understanding of demographic dynamics in preindustrial settings.

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.

Adult sex ratios: causes of variation and implications for animal and human societies.

Converging lines of inquiry from across the social and biological sciences target the adult sex ratio (ASR; the proportion of males in the adult population) as a fundamental population-level determinant of behavior. The ASR, which indicates the relative number of potential mates to competitors in a population, frames the selective arena for competition, mate choice, and social interactions. Here we review a growing literature, focusing on methodological developments that sharpen knowledge of the demographic variables underlying ASR variation, experiments that enhance understanding of the consequences of ASR imbalance across societies, and phylogenetic analyses that provide novel insights into social evolution. We additionally highlight areas where research advances are expected to make accelerating contributions across the social sciences, evolutionary biology, and biodiversity conservation.

The content and structure of reputation domains across human societies: a view from the evolutionary social sciences.

Reputations are an essential feature of human sociality and the evolution of cooperation and group living. Much scholarship has focused on reputations, yet typically on a narrow range of domains (e.g. prosociality and aggressiveness), usually in isolation. Humans can develop reputations, however, from any collective information. We conducted exploratory analyses on the content, distribution and structure of reputation domain diversity across cultures, using the Human Relations Area Files ethnographic database. After coding ethnographic texts on reputations from 153 cultures, we used hierarchical modelling, cluster analysis and text analysis to provide an empirical view of reputation domains across societies. Findings suggest: (i) reputational domains vary cross-culturally, yet reputations for cultural conformity, prosociality, social status and neural capital are widespread; (ii) reputation domains are more variable for males than females; and (iii) particular reputation domains are interrelated, demonstrating a structure consistent with dimensions of human uniqueness. We label these features: cultural group unity, dominance, neural capital, sexuality, social and material success and supernatural healing. We highlight the need for future research on the evolution of cooperation and human sociality to consider a wider range of reputation domains, as well as their social, ecological and gender-specific variability. This article is part of the theme issue 'The language of cooperation: reputation and honest signalling'.

NDVI predicts birth seasonality in historical Baja California Sur, Mexico: adaptive responses to arid ecosystems and the North American Monsoon.

Birth seasonality is a phenomenon whereby populations can be characterized by a single month or season in which births peak. While non-human animal research suggests that seasonal birth-pulses are related to variation in climate and local energy availability, social scientists debate the mechanisms responsible for it in humans. Here we investigate the role of precipitation, temperature, and energy availability on seasonal conception and birth pulses using a historical dataset from the Baja California peninsula - a hot, arid desert that experiences seasonal climatic fluctuations associated with the North American Monsoon. Analyses suggest that 1) local energy availability had a negative relationship with conception pulses; and 2) birth pulses had a positive relationship with local energy availability and a negative relationship with temperature. Taken together, our analyses suggest that women timed conceptions when local energy availability was lowest (challenging expectations of conception rates as simply reflecting ecological influences on female fecundity), so that children were born during the seasonal "green-up" associated with the North American Monsoon. Given our results, we speculate that birth seasonality represents a form of traditional ecological knowledge to improve neonate health and wellbeing.

Was Cinderella just a fairy tale? Survival differences between stepchildren and their half-siblings.

The death of a parent, particularly the mother, is linked to a suite of negative outcomes across the life-course. Compounding concerns for child outcomes are expectations of poor treatment by step-parents after parental remarriage. Indeed, folk tales of step-parental abuse abound cross-culturally and are embedded into stories taught to children. To understand why child outcomes might be sensitive to levels of relatedness within the household, evolutionary-oriented research targets patterning in parental expenditure in ways predicted to maximize inclusive fitness. In particular, parents are expected to prioritize investments in their biological children. However, stepfamilies are only formed after children experience multiple unfortunate events (e.g. parental loss, poverty), blurring causal interpretations between step-parental presence and stepchild outcomes. Moreover, stepchildren have been shown to be integral to household functioning, caring for their half-siblings and stabilizing relationships. These results challenge narrow views of adaptive behaviour; specifically, that step-parents, unlike biological parents, do no stand to reap fitness benefits from the care that they provide to their stepchildren. To evaluate these critiques, we analyse the survival outcomes of stepchildren. We include over 400 000 individuals from across a natural fertility period (1847-1940) in the United States state of Utah and examine the consequences of parental loss and step-parental introduction. Our analyses yield three key results: (i) exposure to maternal loss in childhood is associated with elevated mortality risk, (ii) parental remarriage does not increase the risk of mortality among stepchildren compared to non-stepchildren who too had lost a parent, and (iii) stepchildren enjoy higher survival than their half-siblings within the same family. Ultimately, this work contributes to the increasingly recognized importance of cooperative relationships among non-kin for childcare and household functioning. This article is part of the theme issue 'Multidisciplinary perspectives on social support and maternal-child health'.

Effects of family planning on fertility behaviour across the demographic transition.

The adoption of contraception often coincides with market integration and has transformative effects on fertility behavior. Yet many parents in small-scale societies make decisions about whether and when to adopt family planning in an environment where the payoffs to  have smaller families are uncertain. Here we track the fertility of Maya women across 90 years, spanning the transition from natural to contracepting fertility. We first situate the uncertainty in which fertility decisions are made and model how childbearing behaviors respond. We find that contraception, a key factor in cultural transmission models of fertility decline, initially has little effect on family size as women appear to hedge their bets and adopt fertility control only at the end of their reproductive careers. Family planning is, however, associated with the spread of lower fertility in later cohorts. Distinguishing influences on the origin versus spread of a behaviour provides valuable insight into causal factors shaping individual and normative changes in fertility.

Male survival advantage on the Baja California peninsula.

A consistent finding from contemporary Western societies is that women outlive men. However, what is unclear is whether sex differences in survival are constant across varying socio-ecological conditions. We test the universality of the female survival advantage with mortality data from a nineteenth century population in the Baja California peninsula of Mexico. When examined simply, we find evidence for a male-biased survival advantage. However, results from Cox regression clearly show the importance of age intervals for variable survival patterns by sex. Our key findings are that males: (i) experience significantly lower mortality risk than females during the ages 15-30 (RR = 0.69), (ii) are at a significantly increased risk of dying in the 61+ category (RR = 1.30) and (iii) do not experience significantly different mortality risk at any other age interval (0-14, 31-45, 46-60). We interpret our results to stem from differing intrinsic and extrinsic risk factors for sex-biased mortality across age intervals, highlighting the relevance of a lifecourse approach to the study of survival advantage. Ultimately, our results make clear the need to more broadly consider variability in mortality risk factors across time and place to allow for a clearer understanding of human survival differences.

The role of dispersal and school attendance on reproductive dynamics in small, dispersed populations: Choyeros of Baja California Sur, Mexico.

Individuals from small populations face challenges to initiating reproduction because stochastic demographic processes create local mate scarcity. In response, flexible dispersal patterns that facilitate the movement of individuals across groups have been argued to reduce mate search costs and inbreeding depression. Furthermore, factors that aggregate dispersed peoples, such as rural schools, could lower mate search costs through expansion of mating markets. However, research suggests that dispersal and school attendance are costly to fertility, causing individuals to delay marriage and reproduction. Here, we investigate the role of dispersal and school attendance on marriage and reproductive outcomes using a sample of 54 married couples from four small, dispersed ranching communities in Baja California Sur, Mexico. Our analyses yield three sets of results that challenge conventional expectations. First, we find no evidence that dispersal is associated with later age at marriage or first reproduction for women. For men, dispersal is associated with younger ages of marriage than those who stay in their natal area. Second, in contrast to research suggesting that dispersal decreases inbreeding, we find that female dispersal is associated with an increase in genetic relatedness among marriage partners. This finding suggests that human dispersal promotes female social support from genetic kin in novel locales for raising offspring. Third, counter to typical results on the role of education on reproductive timing, school attendance is associated with younger age at marriage for men and younger age at first birth for women. While we temper causal interpretations and claims of generalizability beyond our study site given our small sample sizes (a feature of small populations), we nonetheless argue that factors like dispersal and school attendance, which are typically associated with delayed reproduction in large population, may actually lower mate search costs in small, dispersed populations with minimal access to labor markets.

Marriage dynamics in old Lower California: ecological constraints and reproductive value in an arid peninsular frontier.

It is commonly expected that natural selection will favor earlier reproduction, yet ecological constraints can force people to delay marriage. Furthermore, humans demonstrate sex-specific preferences in marriage partners - with grooms normally a few years older than their brides; however, the age at which individuals marry can influence the spousal age gap. We investigate factors influencing age at first marriage and age difference at marriage using nineteenth-century historical demographic data from Baja California Sur, Mexico. Analyses suggest ecological constraints affected male, but not female, age at first marriage. Males who migrated from their natal community and who married in communities whose primary economic activity was agriculture experienced delayed age at first marriage. The age at which females first married increased over time causing a reduction in the age gap between spouses. Furthermore, the spousal age gap showed sex-specific effects: women who married early in life were much younger than their husbands, while women who married late in life were older than their husbands, suggesting that variation in female reproductive value influenced mate choice. Males, on the other hand, who married late in life showed a preference for marrying much younger females, indicating preferences for females with high reproductive value.

Sex ratios at birth vary with environmental harshness but not maternal condition.

The sex ratio at birth (SRB) may be patterned by maternal condition and/or environmental stressors. However, despite decades of research, empirical results from across the social and biological sciences are equivocal on this topic. Using longitudinal individual-level data from a US population during the interwar period (1918-1939), inclusive of three distinct eras (Spanish Flu, Roaring '20 s, and the Great Depression), we evaluate predictions from two theoretical frameworks used to study patterning in SRB - (1) 'frail males' and (2) adaptive sex-biased investment theory (Trivers-Willard). The first approach centers on greater male susceptibility to exogenous stressors and argues that offspring survival should be expected to differ between 'good' and 'bad' times. The second approach contends that mothers themselves play a direct role in manipulating offspring SRB, and that those in better condition should invest more in sons. In-line with 'frail male' predictions, we find that boys are less likely to be born during the environmentally challenging times of the Spanish Flu and Great Depression. However, we find no evidence that maternal condition is associated with sex ratios at birth, a result inconsistent with the Trivers-Willard hypothesis.

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.

Adult sex ratios and reproductive strategies: a critical re-examination of sex differences in human and animal societies.

It is increasingly recognized that the relative proportion of potential mates to competitors in a population impacts a range of sex-specific behaviours and in particular mating and reproduction. However, while the adult sex ratio (ASR) has long been recognized as an important link between demography and behaviour, this relationship remains understudied. Here, we introduce the first inter-disciplinary collection of research on the causes and consequences of variation in the ASR in human and animal societies. This important topic is relevant to a wide audience of both social and biological scientists due to the central role that the relative number of males to females in a population plays for the evolution of, and contemporary variation in, sex roles across groups, species and higher taxa. The articles in this theme issue cover research on ASR across a variety of taxa and topics. They offer critical re-evaluations of theoretical foundations within both evolutionary and non-evolutionary fields, and propose innovative methodological approaches, present new empirical examples of behavioural consequences of ASR variation and reveal that the ASR plays a major role in determining population viability, especially in small populations and species with labile sex determination. This introductory paper puts the contributions of the theme issue into a broader context, identifies general trends across the literature and formulates directions for future research.This article is part of the themed issue 'Adult sex ratios and reproductive decisions: a critical re-examination of sex differences in human and animal societies'.

Causes and consequences of adult sex ratio imbalance in a historical U.S. population.

The responsiveness of individuals to partner availability has been well-documented across the literature. However, there is disagreement regarding the direction of the consequences of sex ratio imbalance. Specifically, does an excess of males or females promote male-male mating competition? In an attempt to clarify the role of the adult sex ratio (ASR) on behaviour, here we evaluate both competing and complimentary expectations derived from theory across the social and biological sciences. We use data drawn from a historical, nineteenth century population in North America and target several life-history traits thought to be affected by partner availability: age at first birth, relationship status, completed fertility and longevity. Furthermore, we assess the role of various contributors to a population's ASR. We find that both the contributors to and consequences of sex ratio imbalance vary over time. Our results largely support predictions of greater male pairbond commitment and lesser male mating effort, as well as elevated bargaining power of women in response to female scarcity. After reviewing our findings, and others from across the literature, we highlight the need to adjust predictions in response to ASR imbalance by the: (i) culturally mediated mating arena, (ii) variable role of demographic inputs across time and place, (iii) constraints to behavioural outcomes across populations, and (iv) ability and accuracy of individuals to assess partner availability.This article is part of the themed issue 'Adult sex ratios and reproductive strategies: a critical re-examination of sex differences in human and animal societies'.

Adult sex ratios and partner scarcity among hunter-gatherers: implications for dispersal patterns and the evolution of human sociality.

Small populations are susceptible to high genetic loads and random fluctuations in birth and death rates. While these selective forces can adversely affect their viability, small populations persist across taxa. Here, we investigate the resilience of small groups to demographic uncertainty, and specifically to fluctuations in adult sex ratio (ASR), partner availability and dispersal patterns. Using 25 years of demographic data for two Savannah Pumé groups of South American hunter-gatherers, we show that in small human populations: (i) ASRs fluctuate substantially from year to year, but do not consistently trend in a sex-biased direction; (ii) the primary driver of local variation in partner availability is stochasticity in the sex ratio at maturity; and (iii) dispersal outside of the group is an important behavioural means to mediate locally constrained mating options. To then simulate conditions under which dispersal outside of the local group may have evolved, we develop two mathematical models. Model results predict that if the ASR is biased, the globally rarer sex should disperse. The model's utility is then evaluated by applying our empirical data to this central prediction. The results are consistent with the observed hunter-gatherer pattern of variation in the sex that disperses. Together, these findings offer an alternative explanation to resource provisioning for the evolution of traits central to human sociality (e.g. flexible dispersal, bilocal post-marital residence and cooperation across local groups). We argue that in small populations, looking outside of one's local group is necessary to find a mate and that, motivated by ASR imbalance, the alliances formed to facilitate the movement of partners are an important foundation for the human-typical pattern of network formation across local groups.This article is part of the themed issue 'Adult sex ratios and reproductive decisions: a critical re-examination of sex differences in human and animal societies'.

Marriage Markets and Male Mating Effort: Violence and Crime Are Elevated Where Men Are Rare.

The negative social outcomes in populations with male-biased sex ratios are a growing concern. In general, the expectation is of heightened violence as a result of excess men engaging in antisocial behavior and crime, thereby threatening societal stability. While intuitive, these claims are largely unsupported in the literature. Using mating market theory as our guide, we examine indicators of male mating effort, including (1) violent competition between men (homicide, aggravated assault) and (2) indicators of uncommitted sexual behavior (rape, sex offenses, and prostitution). Our unit of analysis is U.S. county-level data. We find that counties with more men have lower rates of crime and violent behavior. Our findings challenge conventional claims of male excess leading to elevated levels of violence. Instead, in support of mating market predictions, we find that criminal and violent behavior related to male mating effort is least common in male-biased sex ratios. We discuss the implications of our findings for public policy regarding incarceration and criminal behavior.

The evolution of monogamy in response to partner scarcity.

The evolution of monogamy and paternal care in humans is often argued to have resulted from the needs of our expensive offspring. Recent research challenges this claim, however, contending that promiscuous male competitors and the risk of cuckoldry limit the scope for the evolution of male investment. So how did monogamy first evolve? Links between mating strategies and partner availability may offer resolution. While studies of sex roles commonly assume that optimal mating rates for males are higher, fitness payoffs to monogamy and the maintenance of a single partner can be greater when partners are rare. Thus, partner availability is increasingly recognized as a key variable structuring mating behavior. To apply these recent insights to human evolution, we model three male strategies - multiple mating, mate guarding and paternal care - in response to partner availability. Under assumed ancestral human conditions, we find that male mate guarding, rather than paternal care, drives the evolution of monogamy, as it secures a partner and ensures paternity certainty in the face of more promiscuous competitors. Accordingly, we argue that while paternal investment may be common across human societies, current patterns should not be confused with the reason pairing first evolved.