Fertility decline and the changing dynamics of wealth,status and inequality

In the course of demographic transitions (DTs), two large-scale trends become apparent: (i) the broadly positive association between wealth, status and fertility tends to reverse, and (ii) wealth inequalities increase and then temporarily decrease. We argue that these two broad patterns are linked, through a diversification of reproductive strategies that subsequently converge as populations consume more, become less self-sufficient and increasingly depend on education as a route to socio-economic status. We examine these links using data from 22 mid-transition communities in rural Poland. We identify changing relationships between fertility and multiple measures of wealth, status and inequality. Wealth and status generally have opposing effects on fertility, but these associations vary by community. Where farming remains a viable livelihood, reproductive strategies typical of both pre- and post-DT populations coexist. Fertility is lower and less variable in communities with lower wealth inequality, and macro-level patterns in inequality are generally reproduced at the community level. Our results provide a detailed insight into the changing dynamics of wealth, status and inequality that accompany DTs at the community level where peoples'' social and economic interactions typically take place. We find no evidence to suggest that women with the most educational capital gain wealth advantages from reducing fertility, nor that higher educational capital delays the onset of childbearing in this population. Rather, these patterns reflect changing reproductive preferences during a period of profound economic and social change, with implications for our understanding of reproductive and socio-economic inequalities in transitioning populations.     

Understanding variation in human fertility: what can we learn from evolutionary demography?

Decades of research on human fertility has presented a clear picture of how fertility varies, including its dramatic decline over the last two centuries in most parts of the world. Why fertility varies, both between and within populations, is not nearly so well understood. Fertility is a complex phenomenon, partly physiologically and partly behaviourally determined, thus an interdisciplinary approach is required to understand it. Evolutionary demographers have focused on human fertility since the 1980s. The first wave of evolutionary demographic research made major theoretical and empirical advances, investigating variation in fertility primarily in terms of fitness maximization. Research focused particularly on variation within high-fertility populations and small-scale subsistence societies and also yielded a number of hypotheses for why fitness maximization seems to break down as fertility declines during the demographic transition. A second wave of evolutionary demography research on fertility is now underway, paying much more attention to the cultural and psychological mechanisms underpinning fertility. It is also engaging with the complex, multi-causal nature of fertility variation, and with understanding fertility in complex modern and transitioning societies. Here, we summarize the history of evolutionary demographic work on human fertility, describe the current state of the field, and suggest future directions.     

A synthetic biosocial model of fertility transition: Testing the relative contribution of embodied capital theory,changing cultural norms,and women's labor force participation

This article presents a biosocial model of fertility decline, which integrates ecological‐economic and informational‐cultural hypotheses of fertility transition in a unified theoretical framework. The model is then applied to empirical data collected among 500 women from San Borja, Bolivia, a population undergoing fertility transition. Using a combination of event history analysis, multiple regression, and structural equation modeling, we examine the pathways by which education responds to birth cohort, parental education and network ties, and how age at first birth and total fertility, in turn, respond to birth cohort, social network ties, education, expectations about parental investment, work, and contraceptive use. We find that in addition to secular trends in education, respondent's education is associated with the education of parents, the investment she received from them, and the education of older siblings. Total fertility has dropped over time, partly in response to increased education; moreover, the behavior of other women in a woman's social network predicts both initiation of reproduction and total fertility, while expected parental investment in offspring negatively predicts total fertility. Involvement in paid work that is incompatible with childcare is associated with a later age of first reproduction, but not subsequent fertility. Contraceptive use partially mediates the effect of education and birth cohort on total fertility, but is not a mediator of the effect of social network or expected parental investment on total fertility. Overall, the empirical results provide support for a biosocial model of fertility decline, particularly the embodied capital and cultural pathways. Am J Phys Anthropol 154:322–333, 2014. © 2014 Wiley Periodicals, Inc.     

Why do men seek status? Fitness payoffs to dominance and prestige

In many human societies, high male social status associates with higher fertility, but the means by which status increases lifetime fitness have not been systematically investigated. We analyse the pathways by which male status begets reproductive success in a small-scale, Amerindian society. Men who are more likely to win a dyadic physical confrontation, i.e. dominant men, have higher intra-marital fertility for their age, and men with more community-wide influence, i.e. prestigious men, exhibit both higher intra-marital fertility and lower offspring mortality. Both forms of status elicit support from allies and deference from competitors, but high status men are not provisioned more than their peers. Prestigious but not dominant men marry wives who first give birth at earlier ages, which multivariate analysis suggests is the strongest pathway between status and fitness in this population. Furthermore, men are motivated to pursue status because of fitness gains both within and outside of marital unions: dominant and prestigious men have more in-pair surviving offspring as well as more extra-marital affairs.     

The theoretical and political framing of the population factor in development

The silence about population growth in recent decades has hindered the ability of those concerned with ecological change, resource scarcity, health and educational systems, national security, and other global challenges to look with maximum objectivity at the problems they confront. Two central questions about population—(i) is population growth a problem? and (2) what causes fertility decline?—are often intertwined; if people think the second question implies possible coercion, or fear of upsetting cultures, they can be reluctant to talk about the first. The classic and economic theories explaining the demographic transition assume that couples want many children and they make decisions to have a smaller family when some socio-economic change occurs. However, there are numerous anomalies to this explanation. This paper suggests that the societal changes are neither necessary nor sufficient for family size to fall. Many barriers of non-evidence-based restrictive medical rules, cost, misinformation and social traditions exist between women and the fertility regulation methods and correct information they need to manage their family size. When these barriers are reduced, birth rates tend to decline. Many of the barriers reflect a patriarchal desire to control women, which can be largely explained by evolutionary biology. The theoretical explanations of fertility should (i) attach more weight to the many barriers to voluntary fertility regulation, (ii) recognize that a latent desire to control fertility may be far more prevalent among women than previously understood, and (iii) appreciate that women implicitly and rationally make benefit–cost analyses based on the information they have, wanting modern family planning only after they understand it is a safe option. Once it is understood that fertility can be lowered by purely voluntary means, comfort with talking about the population factor in development will rise.     

The cultural evolution of fertility decline

Cultural evolutionists have long been interested in the problem of why fertility declines as populations develop. By outlining plausible mechanistic links between individual decision-making, information flow in populations and competition between groups, models of cultural evolution offer a novel and powerful approach for integrating multiple levels of explanation of fertility transitions. However, only a modest number of models have been published. Their assumptions often differ from those in other evolutionary approaches to social behaviour, but their empirical predictions are often similar. Here I offer the first overview of cultural evolutionary research on demographic transition, critically compare it with approaches taken by other evolutionary researchers, identify gaps and overlaps, and highlight parallel debates in demography. I suggest that researchers divide their labour between three distinct phases of fertility decline—the origin, spread and maintenance of low fertility—each of which may be driven by different causal processes, at different scales, requiring different theoretical and empirical tools. A comparative, multi-level and mechanistic framework is essential for elucidating both the evolved aspects of our psychology that govern reproductive decision-making, and the social, ecological and cultural contingencies that precipitate and sustain fertility decline.     

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.     

The Behavioral Ecology of Family Planning

Family planning is the usual modern route to producing a small family. Can human behavioral ecology provide a framework for understanding family planning behavior? Hillard S. Kaplan (Yearb. Phys. Anthropol. 39:91–135) has proposed a general theory of human parental investment based on the importance of skills development in children. As modern, skills-based, competitive market economies are established, parental investment strategies would be predicted to become oriented toward producing increasingly competitive offspring in a pattern of coordinated investment in their embodied capital—in other words, skills training along with good health to ensure their long-term productivity. Parental embodied capital and resources are also expected to be associated with motivation to produce competitive offspring. The basic parental investment trade-off between quality and quantity should predict greater investment in fewer children and the adoption of family planning behavior. Data on family planning in two ethnic groups in Northeast India (Khasi and Bengali) currently experiencing early-phase transition into modern market economies from very different social and ecological baselines are examined within this analytical framework. The results show a mixture of strategies in conjunction with family planning that involve decreased as well as increased investment in the embodied capital of children among Bengali and a divergence of investments in education and health among Khasi. These mixtures of strategies provide some insight into the motivations to use family planning in the face of economic transition, given differing local cultural and ecological conditions and the opportunity structures they provide.     

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.     

Changing Patterns of Infant Mortality and Maternal Fertility among Pumé Foragers and Horticulturalists

National census data show that the modern demographic transition—the recent trend toward declining mortality and fertility—is well underway in most countries. A different picture emerges when data from small-scale societies in unindustrialized parts of the world are considered. Many of these small-scale societies are also adapting to rapid changes in their subsistence economies. In this article, we examine the relationship between the pace of acculturation, infant mortality, and fertility levels among Pumé foragers and horticulturalists, two related groups of native South Americans. During the earliest stages of acculturation, Pumé horticulturalists experience not only a rapid drop in infant mortality but also a rise in birth rates. An anthropological view of demographic transitions provides important insight into how small-scale societies are affected by exposure to the labor market economy and has practical applications to effective development initiatives and public health policies.     

Social status,immune response and parasitism in males: a meta-analysis

In male vertebrates, two conflicting paradigms—the energetic costs of high dominance rank and the chronic stress of low rank—have been proposed to explain patterns of immune function and parasitism. To date, neither paradigm has provided a complete explanation for status-related differences in male health. Here, we applied meta-analyses to test for correlations between male social status, immune responses and parasitism. We used an ecoimmunological framework, which proposes that males should re-allocate investment in different immune components depending on the costs of dominance or subordination. Spanning 297 analyses, from 77 studies on several vertebrate taxa, we found that most immune responses were similar between subordinate and dominant males, and neither dominant nor subordinate males consistently invested in predictable immune components. However, subordinate males displayed significantly lower delayed-type hypersensitivity and higher levels of some inflammatory cytokines than dominant males, while dominant males exhibited relatively lower immunoglobulin responses than subordinate males. Despite few differences in immunity, dominant males exhibited consistently higher parasitism than subordinate males, including protozoan blood parasites, ectoparasites and gastrointestinal helminths. We discuss our results in the context of the costs of dominance and subordination and advocate future work that measures both parasitism and immune responses in wild systems.     

Social complexity and linguistic diversity in the Austronesian and Bantu population expansions

Reconstructing the rise and fall of social complexity in human societies through time is fundamental for understanding some of the most important transformations in human history. Phylogenetic methods based on language diversity provide a means to reconstruct pre-historic events and model the transition rates of cultural change through time. We model and compare the evolution of social complexity in Austronesian (n = 88) and Bantu (n = 89) societies, two of the world's largest language families with societies representing a wide spectrum of social complexity. Our results show that in both language families, social complexity tends to build and decline in an incremental fashion, while the Austronesian phylogeny provides evidence for additional severe demographic bottlenecks. We suggest that the greater linguistic diversity of the Austronesian language family than Bantu likely follows the different biogeographic structure of the two regions. Cultural evolution in both the Bantu and Austronesian cases was not a simple linear process, but more of a wave-like process closely tied to the demography of expanding populations and the spatial structure of the colonized regions.     

Impact of internal and external factors on prosocial choices in rhesus macaques

While traditional economic models assume that agents are self-interested, humans and most non-human primates are social species. Therefore, many of decisions they make require the integration of information about other social agents. This study asks to what extent information about social status and the social context in which decisions are taken impact on reward-guided decisions in rhesus macaques. We tested 12 monkeys of varying dominance status in several experimental versions of a two-choice task in which reward could be delivered to self only, only another monkey, both the self and another monkey, or neither. Results showed dominant animals were more prone to make prosocial choices than subordinates, but only when the decision was between a reward for self only and a reward for both self and other. If the choice was between a reward for self only and a reward for other only, no animal expressed altruistic behaviour. Finally, prosocial choices were true social decisions as they were strikingly reduced when the social partner was replaced by a non-social object. These results showed that as in humans, rhesus macaques'' social decisions are adaptive and modulated by social status and the cost associated with being prosocial.This article is part of the theme issue ‘Existence and prevalence of economic behaviours among non-human primates’.     

Community-level education accelerates the cultural evolution of fertility decline

Explaining why fertility declines as populations modernize is a profound theoretical challenge. It remains unclear whether the fundamental drivers are economic or cultural in nature. Cultural evolutionary theory suggests that community-level characteristics, for example average education, can alter how low-fertility preferences are transmitted and adopted. These assumptions have not been empirically tested. Here, we show that community-level education accelerates fertility decline in a way that is neither predicted by individual characteristics, nor by the level of economic modernization in a population. In 22 high-fertility communities in Poland, fertility converged on a smaller family size as average education in the community increased—indeed community-level education had a larger impact on fertility decline than did individual education. This convergence was not driven by educational levels being more homogeneous, but by less educated women having fewer children than expected, and more highly educated social networks, when living among more highly educated neighbours. The average level of education in a community may influence the social partners women interact with, both within and beyond their immediate social environments, altering the reproductive norms they are exposed to. Given a critical mass of highly educated women, less educated neighbours may adopt their reproductive behaviour, accelerating the pace of demographic transition. Individual characteristics alone cannot capture these dynamics and studies relying solely on them may systematically underestimate the importance of cultural transmission in driving fertility declines. Our results are inconsistent with a purely individualistic, rational-actor model of fertility decline and suggest that optimization of reproduction is partly driven by cultural dynamics beyond the individual.     

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.     

What if fertility decline is not permanent? The need for an evolutionarily informed approach to understanding low fertility

‘Demographic transition theory’ assumes that fertility decline is irreversible. This commonly held assumption is based on observations of recent and historical reductions in fertility that accompany modernization and declining mortality. The irreversibility assumption, however, is highly suspect from an evolutionary point of view, because demographic traits are at least partially influenced by genetics and are responsive to social and ecological conditions. Nonetheless, an inevitable shift from high mortality and fertility to low mortality and fertility is used as a guiding framework for projecting human population sizes into the future. This paper reviews some theoretical and empirical evidence suggesting that the assumption of irreversibility is ill-founded, at least without considerable development in theory that incorporates evolutionary and ecological processes. We offer general propositions for how fertility could increase in the future, including natural selection on high fertility variants, the difficulty of maintaining universal norms and preferences in a large, diverse and economically differentiated population, and the escalating resource demands of modernization.     

Men in the demographic transition

Women’s fertility is the focus of most demographic analyses, for in most mammals, and in many preindustrial societies, variance in male fertility, while an interesting biological phenomenon, is irrelevant. Yet in monogamous societies, the reproductive ecology of men, as well as that of women, is important is creating reproductive patterns. In nineteenth-century Sweden, the focus of this study, male reproductive ecology responded to resource conditions: richer men had more children than poorer men. Men’s fertility also interacted with local and historical factors in complex ways to have significant impact on population growth. As a result, “the” demographic transition was local, and locally reversible, in Sweden. Results cannot be simply translated from nineteenth-century studies to current attempts to promote fertility decline, because today, male and female resource-fertility curves differ in shape, not only in magnitude. When we translate studies of fertility decline, it is important to study individual fertility and to discern whether, in any particular case, male and female patterns are similar. Bobbi S. Low is Professor of Resource Ecology at the School of Natural Resources and Environment. Her background is in behavioral and evolutionary ecology and her current research interests are human sex differences in risk-taking and resource use.     

Information use and resource competition: an integrative framework

Organisms may reduce uncertainty regarding how best to exploit their environment by collecting information about resource distribution. We develop a model to demonstrate how competition can facilitate or constrain an individual''s ability to use information when acquiring resources. As resource distribution underpins both selection on information use and the strength and nature of competition between individuals, we demonstrate interdependencies between the two that should be common in nature. Individuals in our model can search for resources either personally or by using social information. We explore selection on social information use across a comprehensive range of ecological conditions, generalizing the producer–scrounger framework to a wide diversity of taxa and resources. We show that resource ecology—defined by scarcity, depletion rate and monopolizability—determines patterns of individual differences in social information use. These differences suggest coevolutionary processes linking dominance systems and social information use, with implications for the evolutionary demography of populations.     

Parental investment and the optimization of human family size

Human reproductive behaviour is marked by exceptional variation at the population and individual level. Human behavioural ecologists propose adaptive hypotheses to explain this variation as shifting phenotypic optima in relation to local socioecological niches. Here we review evidence that variation in fertility (offspring number), in both traditional and modern industrialized populations, represents optimization of the life-history trade-off between reproductive rate and parental investment. While a reliance on correlational methods suggests the true costs of sibling resource competition are often poorly estimated, a range of anthropological and demographic studies confirm that parents balance family size against offspring success. Evidence of optimization is less forthcoming. Declines in fertility associated with modernization are particularly difficult to reconcile with adaptive models, because fertility limitation fails to enhance offspring reproductive success. Yet, considering alternative measures, we show that modern low fertility confers many advantages on offspring, which are probably transmitted to future generations. Evidence from populations that have undergone or initiated demographic transition indicate that these rewards to fertility limitation fall selectively on relatively wealthy individuals. The adaptive significance of modern reproductive behaviour remains difficult to evaluate, but may be best understood in response to rising investment costs of rearing socially and economically competitive offspring.     

Social network- and community-level influences on contraceptive use: evidence from rural Poland

The diffusion of ‘modern’ contraceptives—as a proxy for the spread of low-fertility norms—has long interested researchers wishing to understand global fertility decline. A fundamental question is how local cultural norms and other people''s behaviour influence the probability of contraceptive use, independent of women''s socioeconomic and life-history characteristics. However, few studies have combined data at individual, social network and community levels to simultaneously capture multiple levels of influence. Fewer still have tested if the same predictors matter for different contraceptive types. Here, we use new data from 22 high-fertility communities in Poland to compare predictors of the use of (i) any contraceptives—a proxy for the decision to control fertility—with those of (ii) ‘artificial’ contraceptives—a subset of more culturally taboo methods. We find that the contraceptive behaviour of friends and family is more influential than are women''s own characteristics and that community level characteristics additionally influence contraceptive use. Highly educated neighbours accelerate women''s contraceptive use overall, but not their artificial method use. Highly religious neighbours slow women''s artificial method use, but not their contraceptive use overall. Our results highlight different dimensions of sociocultural influence on contraceptive diffusion and suggest that these may be more influential than are individual characteristics. A comparative multilevel framework is needed to understand these dynamics.