The year 2021 marked the fiftieth anniversary of the publication of Robert C. Dunnell's (Free Press, 1971) diminutive yet dense Systematics in prehistory. At the height of the debate between Culture History and New Archaeology, Dunnell's work sought to address a more fundamental issue that was and still is relevant to all branches of prehistoric archaeology, and especially to the study of the Palaeolithic: systematics. Dunnell himself was notorious and controversial, but the importance of his work remains underappreciated. Like other precocious works of that tumultuous time, Systematics in prehistory today remains absent from most course reading lists and gathers dust on library shelves. In this contribution we argue for a greater appreciation of its as yet unfulfilled conceptual and analytical promise. In particular, we briefly chart its somewhat delayed impact via evolutionary archaeology, including how it has also influenced non-Anglophone traditions, especially in South America. The obstinate persistence of classification issues in palaeoanthropology and palaeoarchaeology, we argue, warrants a second look at Dunnell's Systematics.

The form–function conceptual framework, which assumes a strong relationship between the structure of a particular trait and its function, has been crucial for understanding morphological variation and locomotion among extant and fossil species across many disciplines. In biological anthropology, it is the lens through which many important questions and hypotheses have been tackled with respect to relationships between morphology and locomotor kinematics, energetics and performance. However, it is becoming increasingly evident that the morphologies of fossil hominins, apes and humans can confer considerable locomotor diversity and flexibility, and can do so with a range of kinematics depending on soft tissue plasticity and environmental and cultural factors. This complexity is not built into traditional biomechanical or mathematical models of relationships between structure and kinematics or energetics, limiting our interpretation of what bone structure is telling us about behaviour in the past. The nine papers presented in this Special Collection together address some of the challenges that variation in the relationship between form and function pose in evolutionary biomechanics, to better characterise the complexity linking structure and function and to provide tools through which we may begin to incorporate some of this complexity into our functional interpretations.

Ecological and genetic factors have influenced the composition of the human microbiome during our evolutionary history. We analysed the oral microbiota of the Agta, a hunter–gatherer population where some members have adopted an agricultural diet. We show that age is the strongest factor modulating the microbiome, probably through immunosenescence since we identified an increase in the number of species classified as pathogens with age. We also characterised biological and cultural processes generating sexual dimorphism in the oral microbiome. A small subset of oral bacteria is influenced by the host genome, linking host collagen genes to bacterial biofilm formation. Our data also suggest that shifting from a fish/meat diet to a rice-rich diet transforms their microbiome, mirroring the Neolithic transition. All of these factors have implications in the epidemiology of oral diseases. Thus, the human oral microbiome is multifactorial and shaped by various ecological and social factors that modify the oral environment.

Social learning is a critical adaptation for dealing with different forms of variability. Uncertainty is a severe form of variability where the space of possible decisions or probabilities of associated outcomes are unknown. We identified four theoretically important sources of uncertainty: temporal environmental variability; payoff ambiguity; selection-set size; and effective lifespan. When these combine, it is nearly impossible to fully learn about the environment. We develop an evolutionary agent-based model to test how each form of uncertainty affects the evolution of social learning. Agents perform one of several behaviours, modelled as a multi-armed bandit, to acquire payoffs. All agents learn about behavioural payoffs individually through an adaptive behaviour-choice model that uses a softmax decision rule. Use of vertical and oblique payoff-biased social learning evolved to serve as a scaffold for adaptive individual learning – they are not opposite strategies. Different types of uncertainty had varying effects. Temporal environmental variability suppressed social learning, whereas larger selection-set size promoted social learning, even when the environment changed frequently. Payoff ambiguity and lifespan interacted with other uncertainty parameters. This study begins to explain how social learning can predominate despite highly variable real-world environments when effective individual learning helps individuals recover from learning outdated social information.

Cousin marriage, a spousal union between close kin, occurs at high frequencies in many parts of the world. The rates of cousin marriage in humans are concordant with empirical studies that challenge the traditionally held view that reproduction with kin is generally avoided in animals. Similarly, some theoretical models in animal behaviour show that inbreeding avoidance is more constrained than previously thought. Such studies highlight the importance of quantifying the costs and benefits of reproduction among close kin over the whole life-course. Here, we use genealogical data from two human populations with high frequencies of cousin marriage (the Dogon from Mali, and the Ancien Régime nobility from Europe) to estimate these potential costs and benefits. We compare age-specific fertility and survival curves, as well as the projected growth rates, of subpopulations of each marriage type. Fitness costs of cousin marriage are present in terms of reduced child survival (in both populations), while benefits exist as increased fertility for men (in the Dogon) and for women (in the Ancien Régime nobility). We also find some differences in the projected growth rates of lineages as a function of marriage type. Finally, we discuss the trade-offs that might shape marriage decisions in different ecological conditions.

Detecting the forces behind the success or failure of cultural products, such as books or films, remains a challenge. Three such forces are drift, context-biased selection and selection based on content – when things succeed because of their intrinsic appeal. We propose a tool to study content-biased selection in sets of cultural collections – e.g. libraries or movie collections – based on the ‘shortlist effect’: the fact that smaller collections are more selective and more likely to favour highly appealing items over others. We use a model to show that, when the shortlist effect is at work, content-biased cultural selection is associated with greater nestedness in sets of collections. Having established empirically the existence of the shortlist effect, and of content-biased selection, in 28 sets of movie collections, we show that nestedness contributions can be used to estimate to what extent specific movies owe their success to their intrinsic properties. This method can be used in a wide range of datasets to detect the items that owe their success to their intrinsic appeal, as opposed to ‘hidden gems’ or ‘accidental hits’.

We outline the potential for integrating economic and evolutionary approaches to marriage and the family. Our broad argument is that the approaches share a concern for competition. Evolutionary scholars are concerned with the fitness consequences of competition and economists are centrally concerned with the nature of competition: how the allocation of scarce resources is mediated by potentially complex forms of social interaction and conflicts of interest. We illustrate our argument by focusing on conceptual and empirical approaches to a topic of interest to economists and evolutionary scholars: polygynous marriage. In comparing conceptual approaches, we distinguish between those that emphasise the physical environment and those that emphasise the social environment. We discuss some advantages of analysing marriage through the lens of competitive markets, and outline some of the ways that economists analyse the emergence of rules governing the family. In discussing empirical approaches to polygynous marriage, we describe how a concern for informing contemporary policy leads economists to focus on the consequences of polygyny, and in particular we describe some of the ways in which economists attempt to distinguish causal effects from selection effects.

Sexual conflict is a thriving area of animal behaviour research. Yet parallel research in the evolutionary human sciences remains underdeveloped and has become mired by controversy. In this special collection, we aim to invigorate the study of fitness-relevant conflicts between women and men, advocating for three synergistic research priorities. First, we argue that a commitment to diversity is required to innovate the field, achieve ethical research practice, and foster fruitful dialogue with neighbouring social sciences. Accordingly, we have prioritised issues of diversity as editors, aiming to stimulate new connections and perspectives. Second, we call for greater recognition that human sex/gender roles and accompanying conflict behaviours are both subject to natural selection and culturally determined. This motivates our shift in terminology from sexual to gendered conflict when addressing human behaviour, countering stubborn tendencies to essentialise differences between women and men and directing attention to the role of cultural practices, normative sanctions and social learning in structuring conflict battlegrounds. Finally, we draw attention to contemporary policy concerns, including the wellbeing consequences of marriage practices and the gendered implications of market integration. Focus on these themes, combined with attendance to the dangers of ethnocentrism, promises to inform culturally sensitive interventions promoting gender equality worldwide.

Evolutionary perspectives on menopause have focused on explaining why early reproductive cessation in females has emerged and why it is rare throughout the animal kingdom, but less attention has been given to exploring patterns of diversity in age at natural menopause. In this paper, we aim to generate new hypotheses for understanding human patterns of diversity in this trait, defined as age at final menstrual period. To do so, we develop a multilevel, interdisciplinary framework, combining proximate, physiological understandings of ovarian ageing with ultimate, evolutionary perspectives on ageing. We begin by reviewing known patterns of diversity in age at natural menopause in humans, and highlight issues in how menopause is currently defined and measured. Second, we consider together ultimate explanations of menopause timing and proximate understandings of ovarian ageing. We find that ovarian ageing is highly constrained by ageing of the follicle – the somatic structure containing the oocyte – suggesting that menopause timing might be best understood as a by-product of ageing rather than a facultative adaptation. Third, we investigate whether the determinants of somatic senescence also underpin menopause timing. We show that diversity in age at menopause can be, at least partly, explained by the genetic, ecological and life-history determinants of somatic ageing. The public health implications of rethinking menopause as the by-product rather than the catalyst of biological ageing are discussed.

Kinship is a fundamental and universal aspect of the structure of human society. The kinship category of ‘grandparents’ is socially salient, owing to grandparents’ investment in the care of the grandchildren as well as to older generations’ control of wealth and cultural knowledge, but the evolutionary dynamics of grandparent terms has yet to be studied in a phylogenetically explicit context. Here, we present the first phylogenetic comparative study of grandparent terms by investigating 134 languages in Pama–Nyungan, an Australian family of hunter–gatherer languages. We infer that proto-Pama–Nyungan had, with high certainty, four separate terms for grandparents. This state then shifted into either a two-term system that distinguishes the genders of the grandparents or a three-term system that merges the ‘parallel’ grandparents, which could then transition into a different three-term system that merges the ‘cross’ grandparents. We find no support for the co-evolution of these systems with either community marriage organisation or post-marital residence. We find some evidence for the correlation of grandparent and grandchild terms, but no support for the correlation of grandparent and cross-cousin terms, suggesting that grandparents and grandchildren potentially form a single lexical category but that the entire kinship system does not necessarily change synchronously.

Darwin's book on expressions of emotion was one of the first publications to include photographs (Darwin, The expression of the emotions in Man and animals, 1872). The inclusion of expression photographs meant that readers could form their own opinions and could, like Darwin, survey others for their interpretations. As such, the images provided an evidence base and an ‘open source’. Since Darwin, increases in the representativeness and realism of emotional expressions have come from the use of composite images, colour, multiple views and dynamic displays. Research on understanding emotional expressions has been aided by the use of computer graphics to interpolate parametrically between different expressions and to extrapolate exaggerations. This review tracks the developments in how emotions are illustrated and studied and considers where to go next.

The standard theory of rationality posits that agents order preferences according to average utilities associated with different choices. Expected utility theory has repeatedly failed as a predictive theory, as reflected in a growing literature in behavioural economics. Evolutionary theorists have suggested that seemingly irrational behaviours in contemporary contexts may have once served important functions, but existing work linking fitness and choice has not adequately addressed the challenges of constructing an evolutionary theory of decision making. In particular, fitness itself is not a reasonable metric for decision making since its timescale exceeds the lifespan of the decision-maker. Consequently, organisms use proximate systems that work on appropriate timescales and are amenable to feedback and learning. We develop an evolutionary principal–agent model in which individuals utilize a set of proximal choice variables to account for the non-linear dependence of these variables on consumption. While this is insufficient to maximize fitness in the presence of environmental stochasticity, maximum fitness can be achieved by adopting pessimistic probability weightings compatible with the rank-dependent expected utility family of choice models. In particular, pessimistic probability weighting emerges naturally in an evolutionary framework because of extreme intolerance to zeros in multiplicative growth processes.

Standard approaches to cultural evolution focus on the recipients or consumers. This does not take into account the fitness costs incurred in producing the behaviours or artefacts that become cultural, i.e. widespread in a social group. We argue that cultural evolution models should focus on these fitness costs and benefits of cultural production, particularly in the domain of ‘symbolic’ culture. In this approach, cultural products can be considered as a part of the extended phenotype of producers, which can affect the fitness of recipients in a positive way (through cooperation) but also in a detrimental way (through manipulation and exploitation). Taking the producers’ perspective may help explain the specific features of many kinds of cultural products.

The debate on (cultural) group selection regularly suffers from an inclusive fitness overdose. The classical view is that all group selection is kin selection, and that Hamilton's rule works for all models. I claim that not all group selection is kin selection, and that Hamilton's rule does not always get the direction of selection right. More importantly, I will argue that the paper by Smith (2020; Cultural group selection and human cooperation: a conceptual and empirical review. Evolutionary Human Sciences, 2) shows that inclusive fitness is not particularly relevant for much of the empirical evidence relating to the question whether or not cultural group selection shaped human behaviour.

Gender role ideology, i.e. beliefs about how genders should behave, is shaped by social learning. Accordingly, if perceptions about the beliefs of others are inaccurate this may impact trajectories of cultural change. Consistent with this premise, recent studies report evidence of a tendency to overestimate peer support for inequitable gender norms, especially among men, and that correcting apparent ‘norm misperception’ promotes transitions to relatively egalitarian beliefs. However, supporting evidence largely relies on self-report measures vulnerable to social desirability bias. Consequently, observed patterns may reflect researcher measurement error rather than participant misperception. Addressing this shortcoming, we examine men's gender role ideology using both conventional self-reported and a novel wife-reported measure of men's beliefs in an urbanising community in Tanzania. We confirm that participants overestimate peer support for gender inequity. However, the latter measure, which we argue more accurately captures men's true beliefs, implies that this tendency is relatively modest in magnitude and scope. Overestimation was most pronounced among men holding relatively inequitable beliefs, consistent with misperception of peer beliefs reinforcing inequitable norms. Furthermore, older and poorly educated men overestimated peer support for gender inequity the most, suggesting that outdated and limited social information contribute to norm misperception in this context.

From a linguistic standpoint, Proto-Japonic and Proto-Koreanic are assumed to have split off the Transeurasian languages in southern Manchuria. The linguistic idea that Proto-Japonic came earlier than Proto-Koreanic in the chronological scheme means that the Proto-Japonic language first entered the Korean Peninsula, and from there spread to the Japanese archipelago at the beginning of the Yayoi period, around the ninth century BC, while the arrival of Proto-Koreanic in southern Korea is associated with the spread of the rolled rim vessel culture around the fifth century BC. The genealogical sequence of the Pianpu, Mumun and Yayoi cultures, which shared the same pottery production techniques, indicates the spread of Proto-Japonic. On the other hand, migrants moved from Liaodong to the Korean Peninsula and established the rolled rim vessel culture. This population movement was probably due to social and political reasons as the Yan state enlarged its territory eastward. The Proto-Koreanic of the rolled rim vessel culture later spread to the Korean Peninsula and gradually drove out Proto-Japonic, becoming the predecessor of the Koreanic. In this paper, I examine the spread of Proto-Japonic and Proto-Koreanic in Northeast Asia based on archaeological evidence, focusing especially on the genealogy of pottery styles and pottery production techniques.

Models of frequency-dependent social learning posit that individuals respond to the commonality of behaviours without additional variables modifying this. Such strategies bring important trade-offs, e.g. conformity is beneficial when observing people facing the same task but harmful when observing those facing a different task. Instead of rigidly responding to frequencies, however, social learners might modulate their response given additional information. To see, we ran an incentivised experiment where participants played either a game against nature or a coordination game. There were three types of information: (a) choice frequencies in a group of demonstrators; (b) an indication of whether these demonstrators learned in a similar or different environment; and (c) an indication about the reliability of this similarity information. Similarity information was either reliably correct, uninformative or reliably incorrect, where reliably correct and reliably incorrect treatments provided participants with equivalent earning opportunities. Participants adjusted their decision-making to all three types of information. Adjustments, however, were asymmetric, with participants doing especially well when conforming to demonstrators who were reliably similar to them. The overall response, however, was more fluid and complex than this one case. This flexibility should attenuate the trade-offs commonly assumed to shape the evolution of frequency-dependent social learning strategies.

Knowledge and behaviour are transmitted from one individual to another through social learning and eventually disseminated across the population. People often learn useful behaviours socially through selective bias rather than random selection of targets. Prestige bias, or the tendency to selectively imitate prestigious individuals, has been considered an important factor in influencing human behaviour. Although its importance in human society and culture has been recognised, the formulation of prestige bias is less developed than that of other social learning biases. To examine the effects of prestige bias on cultural evolution theoretically, it is imperative to formulate prestige and investigate its basic properties. We reviewed two definitions: one based on first-order cues, such as the demonstrator's appearance and job title, and the other based on second-order cues, such as people's behaviour towards the demonstrator (e.g. people increasingly pay attention to prestigious individuals). This study builds a computational model of prestige bias based on these two definitions and compares the cultural evolutionary dynamics they generate. Our models demonstrate the importance of distinguishing between the two types of formalisation, because they can have different influences on cultural evolution.