Population turnover, behavioral conservatism, and rates of cultural evolution.

Cultural evolution facilitates behavioral adaptation in many species. The pace of cultural evolution can be accelerated by population turnover, where newcomers (immigrants or juvenile recruits) introduce adaptive cultural traits into their new group. However, where newcomers are naïve to the challenges of their new group, population turnover could potentially slow the rate of cultural evolution. Here, we model cultural evolution with population turnover and show that even if turnover results in the replacement of experienced individuals with naïve ones, turnover can still accelerate cultural evolution if (1) the rate of social learning is more than twice as fast as the turnover rate and (b) newcomers are more likely to learn socially than behaviorally conservative existing group members. Although population turnover is a relatively simple factor, it is common to all animal societies, and variation in the turnover rate may potentially play an important role in explaining variation in the occurrence and rates of adaptive cultural evolution across species.

Religious celibacy brings inclusive fitness benefits.

The influence of inclusive fitness interests on the evolution of human institutions remains unclear. Religious celibacy constitutes an especially puzzling institution, often deemed maladaptive. Here, we present sociodemographic data from an agropastoralist Buddhist population in western China, where parents sometimes sent a son to the monastery. We find that men with a monk brother father more children, and grandparents with a monk son have more grandchildren, suggesting that the practice is adaptive. We develop a model of celibacy to elucidate the inclusive fitness costs and benefits associated with this behaviour. We show that a minority of sons being celibate can be favoured if this increases their brothers' reproductive success, but only if the decision is under parental, rather than individual, control. These conditions apply to monks in our study site. Inclusive fitness considerations appear to play a key role in shaping parental preferences to adopt this cultural practice.

How evolutionary behavioural sciences can help us understand behaviour in a pandemic.

The COVID-19 pandemic has brought science into the public eye and to the attention of governments more than ever before. Much of this attention is on work in epidemiology, virology and public health, with most behavioural advice in public health focusing squarely on 'proximate' determinants of behaviour. While epidemiological models are powerful tools to predict the spread of disease when human behaviour is stable, most do not incorporate behavioural change. The evolutionary basis of our preferences and the cultural evolutionary dynamics of our beliefs drive behavioural change, so understanding these evolutionary processes can help inform individual and government decision-making in the face of a pandemic. Lay summary: The COVID-19 pandemic has brought behavioural sciences into the public eye: Without vaccinations, stopping the spread of the virus must rely on behaviour change by limiting contact between people. On the face of it, "stop seeing people" sounds simple. In practice, this is hard. Here we outline how an evolutionary perspective on behaviour change can provide additional insights. Evolutionary theory postulates that our psychology and behaviour did not evolve to maximize our health or that of others. Instead, individuals are expected to act to maximise their inclusive fitness (i.e, spreading our genes) - which can lead to a conflict between behaviours that are in the best interests for the individual, and behaviours that stop the spread of the virus. By examining the ultimate explanations of behaviour related to pandemic-management (such as behavioural compliance and social distancing), we conclude that "good of the group" arguments and "one size fits all" policies are unlikely to encourage behaviour change over the long-term. Sustained behaviour change to keep pandemics at bay is much more likely to emerge from environmental change, so governments and policy makers may need to facilitate significant social change - such as improving life experiences for disadvantaged groups.

Modelling cultural selection on biological fitness to integrate social transmission and adaptive explanations for human behaviour.

One of the difficulties with cultural group selection theory highlighted in the review by Smith (2020, Evol. Hum. Sci., 2, e7) is its inability to separate the evolutionary effects of selection of cultural traits based on biological fitness (Cultural Selection 1) from the effects of selection based on cultural fitness (Cultural Selection 2). Confusing these two processes can hinder the integration of adaptive explanations for human behaviour, which focus on biological fitness, and cultural evolution explanations, which often focus on social transmission. Recent empirical work is starting to bridge this gap, but progress in mathematical modelling has been considerably slower. Here, I suggest that modellers can contribute to achieving this integration by further developing models of Cultural Selection 1, where behaviours are influenced by culturally inherited traits selected on the basis of their effects on biological fitness. These models should build on existing social evolution theory methods and replace genetic relatedness with cultural relatedness, that is the probability that two individuals share a cultural variant.

The demography of human warfare can drive sex differences in altruism.

Recent years have seen great interest in the suggestion that between-group aggression and within-group altruism have coevolved. However, these efforts have neglected the possibility that warfare - via its impact on demography - might influence human social behaviours more widely, not just those directly connected to success in war. Moreover, the potential for sex differences in the demography of warfare to translate into sex differences in social behaviour more generally has remained unexplored. Here, we develop a kin-selection model of altruism performed by men and women for the benefit of their groupmates in a population experiencing intergroup conflict. We find that warfare can promote altruistic, helping behaviours as the additional reproductive opportunities winners obtain in defeated groups decrease harmful competition between kin. Furthermore, we find that sex can be a crucial modulator of altruism, with there being a tendency for the sex that competes more intensely with relatives to behave more altruistically and for the sex that competes more intensely with non-relatives in defeated groups to receive more altruism. In addition, there is also a tendency for the less-dispersing sex to both give and receive more altruism. We discuss implications for our understanding of observed sex differences in cooperation in human societies.

Why war is a man's game.

Interest in the evolutionary origins and drivers of warfare in ancient and contemporary small-scale human societies has greatly increased in the last decade, and has been particularly spurred by exciting archaeological discoveries that suggest our ancestors led more violent lives than previously documented. However, the striking observation that warfare is an almost-exclusively male activity remains unexplained. Three general hypotheses have been proposed, concerning greater male effectiveness in warfare, lower male costs, and patrilocality. But while each of these factors might explain why warfare is more common in men, they do not convincingly explain why women almost never participate. Here, we develop a mathematical model to formally assess these hypotheses. Surprisingly, we find that exclusively male warfare may evolve even in the absence of any such sex differences, though sex biases in these parameters can make this evolutionary outcome more likely. The qualitative observation that participation in warfare is almost exclusive to one sex is ultimately explained by the fundamentally sex-specific nature of Darwinian competition-in fitness terms, men compete with men and women with women. These results reveal a potentially key role for ancestral conditions in shaping our species' patterns of sexual division of labour and violence-related adaptations and behavioural disorders.

Intrafamily and intragenomic conflicts in human warfare.

Recent years have seen an explosion of multidisciplinary interest in ancient human warfare. Theory has emphasized a key role for kin-selected cooperation, modulated by sex-specific demography, in explaining intergroup violence. However, conflicts of interest remain a relatively underexplored factor in the evolutionary-ecological study of warfare, with little consideration given to which parties influence the decision to go to war and how their motivations may differ. We develop a mathematical model to investigate the interplay between sex-specific demography and human warfare, showing that: the ecology of warfare drives the evolution of sex-biased dispersal; sex-biased dispersal modulates intrafamily and intragenomic conflicts in relation to warfare; intragenomic conflict drives parent-of-origin-specific patterns of gene expression-i.e. 'genomic imprinting'-in relation to warfare phenotypes; and an ecological perspective of conflicts at the levels of the gene, individual, and social group yields novel predictions as to pathologies associated with mutations and epimutations at loci underpinning human violence.