Resources and the life course: Patterns through the demographic transition

In most mammals, and in the majority of traditional human societies for which data exist, status, power, or resource control correlates with lifetime reproductive success; male and female patterns differ. Because such correlations are often argued to have disappeared in human societies during the demographic transition of the nineteenth century, we analyzed wealth and lifetime reproductive success in a nineteenth-century Swedish population in four economically diverse parishes, subsuming geographic and temporal variation. Children of both sexes born to poorer parents were more likely than richer children to die or emigrate before reaching maturity. Poorer men, and women whose fathers were poorer, were less likely to marry in the parish than others, largely as a result of differential mortality and migration. Of all adults of both sexes who remained in their home parish and thus generated complete lifetime records, richer individuals had greater lifetime fertility and more children alive at age ten, than others. The age-specific fertility of richer women rises slightly sooner, and reaches a higher peak, than that of poorer women. These patterns persisted throughout the period of the sample (1824–1896). Thus, wealth appears, even during the demographic transition in an egalitarian society, to have influenced lifetime reproductive success positively.     

Ecological demography: A synthetic focus in evolutionary anthropology

The interests of evolutionary anthropologists, behavioral ecologists, and demographers converge on the ecology of human fertility. Ecological conditions influence the optimum pattern of maternal effort. Patterns of abortion, neglect, and infanticide vary with mothers' ability to invest in their children and children's ability to use that investment. As in most other mammals, the ecology of human fertility varies between the sexes: status and resource control are important for males, whereas reproductive value is crucial for females. In pre-industrial societies, and even in monogamous societies in demographic transition, wealthy men had more children than did poorer men. This correlation, often assumed to have disappeared, persists today, with richer men still having more sexual access than others. Sex differences in the ecology of fertility mean that sex of the offspring, as well as birth order, influences parental investment. Because individual fertility varies with environment, it is not surprising that “natural” (uncontrolled) fertility varies across societies or that demographic transitions proceed locally, with occasional reverses, as individuals strive to maximize their lifetime reproductive success in changing, competitive, conditions.     

Women,resources, and dispersal in nineteenth-century Sweden

In a recent study, female dispersal in nineteenth-century Sweden has been found to correlate negatively with access to resources: women with limited access to local resources tended to migrate more frequently. In this paper I review the literature to explore whether this observed correlation was derived from a relationship in which a woman’s limited access to resources worsened her position in the marriage market and led to migration, as a strategy to improve resources and this position. Many studies within a variety of disciplines indicate that a woman’s propensity to disperse from her parish of birth variedinversely with her propensity to inherit resources. My review of the literature suggests that the less likely a woman was to inherit resources, the lower her probability of marriage, the later her expected age at marriage, and the earlier she left home, presumably to improve her resource base for marriage. This work was funded in part by the Population and Environment Dynamics Program and by the Evolution and Human Behavior Program, University of Michigan. Alice L. Clarke is a doctoral candidate in the School of Natural Resources and Environment, University of Michigan, Ann Arbor. Her dissertation research examines the ecology of human dispersal in nineteenth-century Sweden. Her broader research interests include the ecological basis of human demography and resource use.     

How Grandparents Matter

Low birth rates in developed societies reflect women’s difficulties in combining work and motherhood. While demographic research has focused on the role of formal childcare in easing this dilemma, evolutionary theory points to the importance of kin. The cooperative breeding hypothesis states that the wider kin group has facilitated women’s reproduction during our evolutionary history. This mechanism has been demonstrated in pre-industrial societies, but there is no direct evidence of beneficial effects of kin’s support on parents’ reproduction in modern societies. Using three-generation longitudinal data anchored in a sample of grandparents aged 55 and over in 1992 in the Netherlands, we show that childcare support from grandparents increases the probability that parents have additional children in the next 8 to 10 years. Grandparental childcare provided to a nephew or niece of childless children did not significantly increase the probability that those children started a family. These results suggest that childcare support by grandparents can enhance their children’s reproductive success in modern societies and is an important factor in people’s fertility decisions, along with the availability of formal childcare.     

Status,reproductive success,and marrying polygynously

A study of female reproductive histories from nineteenth-century Utah shows that although women who married polygynously had fewer children, their number of grandchildren was equal to that of women who married monogamously. Women who chose to marry high-status men polygynously traded decreased fertility for enhanced reproductive performance of offspring. High status can be associated with low fertility and yet still be consistent with fitness optimization. These results suggest how female reproductive decisions influence social structure and challenge previous assumptions concerning proximate measures of fitness.     

Why men matter: mating patterns drive evolution of human lifespan

Evolutionary theory predicts that senescence, a decline in survival rates with age, is the consequence of stronger selection on alleles that affect fertility or mortality earlier rather than later in life. Hamilton quantified this argument by showing that a rare mutation reducing survival is opposed by a selective force that declines with age over reproductive life. He used a female-only demographic model, predicting that female menopause at age ca. 50 yrs should be followed by a sharp increase in mortality, a "wall of death." Human lives obviously do not display such a wall. Explanations of the evolution of lifespan beyond the age of female menopause have proven difficult to describe as explicit genetic models. Here we argue that the inclusion of males and mating patterns extends Hamilton's theory and predicts the pattern of human senescence. We analyze a general two-sex model to show that selection favors survival for as long as men reproduce. Male fertility can only result from matings with fertile females, and we present a range of data showing that males much older than 50 yrs have substantial realized fertility through matings with younger females, a pattern that was likely typical among early humans. Thus old-age male fertility provides a selective force against autosomal deleterious mutations at ages far past female menopause with no sharp upper age limit, eliminating the wall of death. Our findings illustrate the evolutionary importance of males and mating preferences, and show that one-sex demographic models are insufficient to describe the forces that shape human senescence.     

Preferential parental investment in daughters over sons

Female-biased parental investment is unusual but not unknown in human societies. Relevant explanatory models include Fisher’s principle, the Trivers-Willard model, local mate and resource competition and enhancement, and economic rational actor models. Possible evidence of female-biased parental investment includes sex ratios, mortality rates, parents’ stated preferences for offspring of one sex, and direct and indirect measurements of actual parental behavior. Possible examples of female-biased parental investment include the Mukogodo of Kenya, the Ifalukese of Micronesia, the Cheyenne of North America, the Herero of southern Africa, the Kanjar of south Asia, the Mundugumor of New Guinea, contemporary North America, and historical Germany, Portugal, and the United States. Lee Cronk is Assistant Professor of Anthropology at Texas A&M University, College Station, Texas. His main research interests are in human behavioral ecology, reproductive strategies, and East African hunter-gatherers and pastoralists.     

The evolution of magnanimity

Conspicuous consumption associated with status reinforcement behavior can be explained in terms of costly signaling, or strategic handicap theory, first articulated by Zahavi and later formalized by Grafen. A theory is introduced which suggests that the evolutionary raison d’être of status reinforcement behavior lies not only in its effects on lifetime reproductive success, but in its positive effects on the probability of survival through infrequent, unpredictable demographic bottlenecks. Under some circumstances, such “wasteful” displays may take the form of displays of altruistic behavior and generosity on the part of high status individuals, in that is signals the ability to bear the short-term costs of being generous or “cooperative,” while at the same time reinforcing the long-term benefits of higher status. James L. Boone is an associate professor in the Department of Anthropology at the University of New Mexico, where he carries out research in behavioral ecology and the archaeology of complex societies. His current interests are in the evolution of social status reinforcement behavior and variation in patterns of conspicuous consumption in human history.     

The life histories of American stepfathers in evolutionary perspective

This paper presents an analysis of the characteristics of men who become stepfathers, and their subsequent fertility patterns and lifetime reproductive success. Because women who already have children are ranked lower in the marriage market than women without children, men who marry women with children (e.g., stepfathers) are likely to have lower rankings in the marriage market as well. Using retrospective fertility and marital histories from the Panel Study of Income Dynamics (PSID), I show that men who become stepfathers have lower levels of education, less income, and are more likely to have been divorced before and to already have children, all characteristics that lower their rankings in the marriage market. Men with one or two stepchildren are just as likely to have children within a marriage as non-stepfathers, although men with three stepchildren show decreased fertility. Among men age 45 and older, stepfathers have lower lifetime fertility than non-stepfathers, although the difference disappears when men’s age at first marriage is controlled for. Additionally, stepfathers have significantly higher fertility than men who never marry. The results suggest that some men become stepfathers to procure mates and fertility benefits that they would otherwise have been unlikely to obtain; for these men, raising other men’s children serves as a form of mating effort. Preliminary versions of this paper were presented at the Evolution and Human Adaptation Program at the University of Michigan, and the Human Behavior and Evolution Society’s annual meeting at Amherst. Kermyt G. Anderson received his Ph.D. in anthropology from the University of New Mexico in 1999. He is currently a Mellon Postdoctoral Research Fellow at the Population Studies Center of the Institute for Social Research at the University of Michigan. His current research examines the relationship between family structure, parental investment, and children’s educational and employment outcomes in South Africa.     

Mate choice in modern societies

Most research on mate choice in modern societies is based on data that may or may not reflect actual mating behavior (e.g., stated preferences, personal advertisements). In the present study, real-life matings were reported by a large representative sample of men and women (N = 1,133). These data were used to test an evolutionary model in which mate choice is hypothesized to depend on resources potentially contributed to reproduction by each sex. Consistent with the model, it was found that (a) men (but not women) of higher social status acquire more mating partners, suggesting that male status is an important criterion in female choice; (b) women’s (but not men’s) number of partners decreases linearly with age, suggesting that female reproductive potential is an important criterion in male choice; and (c) women (but not men) display a significant relationship between marital dissolution and promiscuity, suggesting that female sexual exclusivity is an important criterion in male choice. These results are discussed in relation to understanding mate choice mechanisms from behavioral data. Daniel Pérusse is an assistant professor in the Department of Anthropology at the Université de Montréal. His research interests include the evolutionary biology of human social and reproductive behavior, sexual selection theory, and biocultural evolution. His current research bears on human socialization processes and psychosocial development from an evolutionary and behavior-genetic perspective. Recent publications include “Cultural and Reproductive Success in Industrial Societies: Testing the Relationship at the Proximate and Ultimate Levels” (Behavioral and Brain Sciences 16(2):267–322, 1993) and “Human Parental Behavior: Evidence for Genetic Influence and Potential Implications for Gene-Culture Theory” with M. C. Neale, A. Heath, and L. J. Eaves (Behavior Genetics, in press).     

Mate Choice and the Origin of Menopause

Human menopause is an unsolved evolutionary puzzle, and relationships among the factors that produced it remain understood poorly. Classic theory, involving a one-sex (female) model of human demography, suggests that genes imparting deleterious effects on post-reproductive survival will accumulate. Thus, a ‘death barrier’ should emerge beyond the maximum age for female reproduction. Under this scenario, few women would experience menopause (decreased fertility with continued survival) because few would survive much longer than they reproduced. However, no death barrier is observed in human populations. Subsequent theoretical research has shown that two-sex models, including male fertility at older ages, avoid the death barrier. Here we use a stochastic, two-sex computational model implemented by computer simulation to show how male mating preference for younger females could lead to the accumulation of mutations deleterious to female fertility and thus produce a menopausal period. Our model requires neither the initial assumption of a decline in older female fertility nor the effects of inclusive fitness through which older, non-reproducing women assist in the reproductive efforts of younger women. Our model helps to explain why such effects, observed in many societies, may be insufficient factors in elucidating the origin of menopause.     

Demographic studies enhance the understanding of evolutionarily (mal)adaptive behaviors and phenomena in humans: a review on fertility decline and an integrated model

Recently, statistical analyses of demographic datasets have come to play an important role for studies into the evolution of human life history. In the first part of this paper, I highlight fertility decline, an evolutionarily paradoxical phenomenon in terms of fitness maximization. Then, I conduct a literature review regarding the effects of socioeconomic status on the number of offspring, especially in modern developed, (post-)industrial, and low-fertility societies. Although a non-positive relationship between them has often been recognized as a general feature of fertility decline, there actually exists a great deal of variation. Based on the review, I discuss the association between socioeconomic success and reproductive success, and tackle an evolutionary question as to why people seek higher socioeconomic success that would not directly lead to higher reproductive success. It has been suggested that, in modern competitive environments, parents should set a higher value on their investment in children, and aim to have a smaller number of high-quality children. Also, parents would maintain higher socioeconomic status for themselves so as to provide high-levels of investment in their children. In the second part, I broadly consider seemingly evolutionarily (mal)adaptive outcomes besides fertility decline, including child abuse, menopause, and suicide. The integration of the major three approaches to human behavioral and psychological research (behavioral ecology, evolutionary psychology, and cultural evolution) could lead to a deeper understanding. I provide a model for the integrated approach. Rich empirical evidence accumulated in demographic studies, especially longitudinal and cross-cultural resources, can assist to develop a theoretical framework.     

Life Course Perspectives on Women's Autonomy and Health Outcomes

Gender inequality leads to negative demographic consequences in many societies. Patterns of household formation and inheritance strongly influence these consequences. Peasant societies of preindustrial northern Europe emphasized the conjugal bond, while intergenerational bonds were weak. The reverse is true in contemporary northern India. As a result, greater potential exists there for marginalizing women. The convergence of low autonomy due to youth as well as gender means that women's autonomy is at its lowest point during the peak of childbearing years. This has considerable implications for demographic and health outcomes in terms of poorer child survival, slower fertility decline, and poorer reproductive health.     

Behavioral ecology of conservation in traditional societies

A common exhortation by conservationists suggests that we can solve ecological problems by returning to the attitudes of traditional societies: reverence for resources, and willingness to assume short-term individual costs for long-term, group-beneficial sustainable management. This paper uses the 186-society Standard Cross-Cultural Sample to examine resource attitudes and practices. Two main findings emerge: (1) resource practices are ecologically driven and do not appear to correlate with attitude (including sacred prohibition) and (2) the low ecological impact of many traditional societies results not from conscious conservation efforts, but from various combinations of low population density, inefficient extraction technology, and lack of profitable markets for extracted resources. Professor Emilio Moran, Director of the Anthropological Center for Training and Research on Global Environmental Change, Indiana University, kindly shared Marcos Terena’s words. Bobbi Low is Professor of Resource Ecology, School of Natural Resources and Environment, University of Michigan. Her research interests include sex differences in resource use, patterns in resource use and reproduction/family formation, and demographic transition patterns.     

Female reproductive behaviour,ovarian hormones and vaginal cytology of the induced ovulator, <Emphasis Type="Italic">Ctenomys talarum</Emphasis>

In the present study, we evaluated whether reproductive condition affects female reproductive behaviour in the induced ovulator Ctenomys talarum. We also explored the effect of the interaction with a male on the reproductive condition of females. To evaluate this, we arranged mating trials and evaluated female reproductive behaviour. Reproductive status of females was evaluated using a combined approach of vaginal smears, urinary progesterone and oestradiol, and ovarian histology. Behaviours denoting attraction (‘male sniff’ and ‘mount attempts’) and mutual courtship behaviours (‘spin’ and copula) were correlated with vaginal cytology before and oestradiol and progesterone levels in urine 12 h after male–female encounter. After 24 h of the interaction, oestradiol levels and vaginal epithelization increased while progesterone levels decreased in soliciting females. C. talarum females’ reproductive behaviour was related to its physiological reproductive state and vaginal cytology. The kind of male interaction, whether couples copulated or remained indifferent affected the later status of females. Females are induced ovulators by mating but male presence and interaction also affected other components of their reproductive physiology such as ovarian hormones and vaginal cytology.     

Does observed fertility maximize fitness among New Mexican men?

Our objective is to test an optimality model of human fertility that specifies the behavioral requirements for fitness maximization in order (a) to determine whether current behavior does maximize fitness and, if not, (b) to use the specific nature of the behavioral deviations from fitness maximization towards the development of models of evolved proximate mechanisms that may have maximized fitness in the past but lead to deviations under present conditions. To test the model we use data from a representative sample of 7,107 men living in Albuquerque, New Mexico, between 1990 and 1993. The model we test proposes that low fertility in modern settings maximizes number of grandchildren as a result of a trade-off between parental fertility and next generation fertility. Results do not show the optimization, although the data do reveal a trade-off between parental fertility and offspring education and income. We propose that two characteristics of modern economies have led to a period of sustained fertility reduction and to a corresponding lack of association between income and fertility. The first is the direct link between costs of investment and wage rates due to the forces of supply and demand for labor in competitive economies. The second is the increasing emphasis on cumulative knowledge, skills, and technologies in the production of resources. Together they produce historically novel conditions. These two features of modern economies may interact with evolved psychological and physiological mechanisms governing fertility and parental investment to produce behavior that maximizes the economic productivity of lineages at the expense of fitness. If cognitive processes evolved to track diminishing returns to parental investment and if physiological processes evolved to regulate fertility in response to nutritional state and patterns of breast feeding, we might expect non-adaptive responses when returns from parental investment do not diminish until extremely high levels are reached. With high economic payoffs from parental investment, people have begun to exercise cognitive regulation of fertility through contraception and family planning practices. Those cognitive processes maynot have evolved to handle fitness trade-offs between fertility and parental investment. A preliminary presentation of this data was published in R. I. M. Dunbar, ed.,Human Reproduction Decisions: Biological and Social Perspectives. New York: St. Martin’s Press, 1995. Support for the research project, “Male Fertility and Parenting in New Mexico,” began with two seed grants from the University of New Mexico’s Biomedical Research Grants Program, 1988 and 1989, and one from the University of New Mexico Research Allocations Committee, 1988. Further seed money as well as interim funding came from the William T. Grant Foundation (#89130589 and #91130501). The major support for the project came from the National Science Foundation from 1990 to 1993 (#BNS-9011723 and #DBS-911552). Both National Science Foundation grants included Research Experience for Undergraduates supplements. Hillard S. Kaplan is an Associate Professor of Anthropology at the University of New Mexico. His earlier research and publications focused on food sharing, time allocation, parental investment, and reproductive strategies among Ache hunter-gatherers in Paraguay, Machiguenga and Piro forager-horticulturalists in Peru, and villagers of several ethnicities in Botswana. New research and theory concern fertility, parental investment, and mating strategies in developed and developing nations. This research formulates a new theory of reproductive decision-making and the demographic transition, integrating human capital and parental investment theory in a synthesis of economic and evolutionary approaches. Jane B. Lancaster is a Professor of Anthropology at the University of New Mexico. Her research and publications are on human reproductive biology and behavior, especially human parental investment; women’s reproductive biology of pregnancy, lactation, and child-spacing; and male fertility and investment in children. Current research with Hillard S. Kaplan is on male life history strategies among a large sample of men in New Mexico. She has coedited three books on human parental investment:School-Age Pregnancy and Parenthood (with B. Hamburg),Parenting across the Life Span (with J. Altmann, A. Rossi, and L. Sherrod), andOffspring Abuse and Neglect (with R. Gelles). She is scientific editor of a quarterly journal,Human Nature: An Interdisciplinary, Biosocial Perspective published by Aldine de Gruyter. She is also a council member of the newly formed Human Behavior and Evolution Society. John A. Bock is Andrew W. Mellon Post-Doctoral Fellow in Epidemiology and Population Health at the National Centre for Epidemiology and Population Health, The Australian National University. His research focuses on the allocation of parental investment and the determinants of children’s activities, integrating aspects of economic and evolutionary theory. He has ongoing field research with Bantu and Bushmen agro-pastoralists and forager-horticulturalists in the Okavango Delta, Botswana. He is also collaborating with Lancaster and Kaplan on the determinants of progeny distribution and homosexuality among New Mexican men. Sara E. Johnson is a Ph.D. candidate at the University of New Mexico. Her major research trajectory focuses on trade-offs in life history characters. Her research experience includes participation in a study of variation in growth and development among children in a multi-ethnic community in the Okavango Delta, Botswana, in addition to her dissertation work on individual variation in growth and mortality among juvenile baboons. She is collaborating with Lancaster and Kaplan on the association between survival and fertility among Albuquerque men.     

Human population growth and the demographic transition

The world and most regions and countries are experiencing unprecedentedly rapid demographic change. The most obvious example of this change is the huge expansion of human numbers: four billion have been added since 1950. Projections for the next half century expect a highly divergent world, with stagnation or potential decline in parts of the developed world and continued rapid growth in the least developed regions. Other demographic processes are also undergoing extraordinary change: women''s fertility has dropped rapidly and life expectancy has risen to new highs. Past trends in fertility and mortality have led to very young populations in high fertility countries in the developing world and to increasingly older populations in the developed world. Contemporary societies are now at very different stages of their demographic transitions. This paper summarizes key trends in population size, fertility and mortality, and age structures during these transitions. The focus is on the century from 1950 to 2050, which covers the period of most rapid global demographic transformation.     

Variation in Male Reproductive Longevity across Traditional Societies

Most accounts of human life history propose that women have short reproductive spans relative to their adult lifespans, while men not only remain fertile but carry on reproducing until late life. Here we argue that studies have overlooked evidence for variation in male reproductive ageing across human populations. We apply a Bayesian approach to census data from Agta hunter-gatherers and Gambian farmers to show that long post-reproductive lifespans characterise not only women but also males in some traditional human populations. We calculate three indices of reproductive ageing in men (oldest age at reproduction, male late-life reproduction, and post-reproductive representation) and identify a continuum of male reproductive longevity across eight traditional societies ranging from !Kung, Hadza and Agta hunter-gatherers exhibiting low levels of polygyny, early age at last reproduction and long post-reproductive lifespans, to male Gambian agriculturalists and Turkana pastoralists showing higher levels of polygyny, late-life reproduction and shorter post-reproductive lifespans. We conclude that the uniquely human detachment between rates of somatic senescence and reproductive decline, and the existence of post-reproductive lifespans, are features of both male and female life histories, and therefore not exclusive consequences of menopause.     

Middle Eastern masculinities in the age of new reproductive technologies: male infertility and stigma in Egypt and Lebanon

Worldwide, male infertility contributes to more than half of all cases of childlessness; yet, it is a reproductive health problem that is poorly studied and understood. This article examines the problem of male infertility in two Middle Eastern locales, Cairo, Egypt, and Beirut, Lebanon, where men may be at increased risk of male infertility because of environmental and behavioral factors. It is argued that male infertility may be particularly problematic for Middle Eastern men in their pronatalist societies; there, both virility and fertility are typically tied to manhood. Thus, male infertility is a potentially emasculating condition, surrounded by secrecy and stigma. Furthermore, the new reproductive technology called intracytoplasmic sperm injection (ICSI), designed specifically to overcome male infertility, may paradoxically create additional layers of stigma and secrecy, due to the complex moral and marital dilemmas associated with Islamic restrictions on third-party donation of gametes.     

Status competition,inequality, and fertility: implications for the demographic transition

The role that social status plays in small-scale societies suggests that status may be important for understanding the evolution of human fertility decisions, and for understanding how such decisions play out in modern contexts. This paper explores whether modelling competition for status—in the sense of relative rank within a society—can help shed light on fertility decline and the demographic transition. We develop a model of how levels of inequality and status competition affect optimal investment by parents in the embodied capital (health, strength, and skills) and social status of offspring, focusing on feedbacks between individual decisions and socio-ecological conditions. We find that conditions similar to those in demographic transition societies yield increased investment in both embodied capital and social status, generating substantial decreases in fertility, particularly under conditions of high inequality and intense status competition. We suggest that a complete explanation for both fertility variation in small-scale societies and modern fertility decline will take into account the effects of status competition and inequality.