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.

Humans have adapted to an immense array of environments by accumulating culturally transmitted knowledge and skills. Adaptive culture can accumulate either via more distinct cultural traits or via improvements of existing cultural traits. The kind of culture that accumulates depends on, and coevolves with, the social structure of societies. Here, we show that the coevolution of learning networks and cumulative culture results in two distinct pathways to cultural adaptation: highly connected populations with high proficiency but low trait diversity vs. sparsely connected populations with low proficiency but higher trait diversity. Importantly, we show there is a conflict between group-level payoffs, which are maximised in highly connected groups that attain high proficiency, and individual level selection, which favours disconnection. This conflict emerges from the interaction of social learning with population structure and causes populations to cycle between the two cultural and network states. The same conflict creates a paradox where increasing innovation rate lowers group payoffs. Finally, we explore how populations navigate these two pathways in environments where payoffs differ among traits and can change over time, showing that high proficiency is favoured when payoffs are stable and vary strongly between traits, while frequent changes in trait payoffs favour more trait diversity. Our results illustrate the complex interplay between networks, learning and the environment, and so inform our understanding of human social evolution.

Among vertebrates, allomothering (non-maternal care) is classified as cooperative breeding (help from sexually mature non-breeders, usually close relatives) or communal breeding (shared care between multiple breeders who are not necessarily related). Humans have been described with both labels, most frequently as cooperative breeders. However, few studies have quantified the relative contributions of allomothers according to whether they are (a) sexually mature and reproductively active and (b) related or unrelated. We constructed close-proximity networks of Agta and BaYaka hunter–gatherers. We used portable remote-sensing devices to quantify the proportion of time children under the age of 4 spent in close proximity to different categories of potential allomother. Both related and unrelated, and reproductively active and inactive, campmates had substantial involvement in children's close-proximity networks. Unrelated campmates, siblings and subadults were the most involved in both populations, whereas the involvement of fathers and grandmothers was the most variable between the two populations. Finally, the involvement of sexually mature, reproductively inactive adults was low. Where possible, we compared our findings with studies of other hunter–gatherer societies, and observed numerous consistent trends. Based on our results we discuss why hunter–gatherer allomothering cannot be fully characterised as cooperative or communal breeding.

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.

Mate preferences and mating-related behaviours are hypothesised to change over the menstrual cycle to increase reproductive fitness. Recent large-scale studies suggest that previously reported hormone-linked behavioural changes are not robust. The proposal that women's preference for associating with male kin is down-regulated during the ovulatory (high-fertility) phase of the menstrual cycle to reduce inbreeding has not been tested in large samples. Consequently, we investigated the relationship between longitudinal changes in women's steroid hormone levels and their perceptions of faces experimentally manipulated to possess kinship cues (Study 1). Women viewed faces displaying kinship cues as more attractive and trustworthy, but this effect was not related to hormonal proxies of conception risk. Study 2 employed a daily diary approach and found no evidence that women spent less time with kin generally or with male kin specifically during the fertile phase of the menstrual cycle. Thus, neither study found evidence that inbreeding avoidance is up-regulated during the ovulatory phase of the menstrual cycle.

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.

Building cooperative communities is a crucial problem for human societies. Much research suggests that cooperation is facilitated by knowing who the cooperators and defectors are, and being able to respond accordingly. As such, anonymous games are thought to hinder cooperation. Here, we show that this conclusion is altered dramatically in the presence of conditional cooperation norms and heterogeneous beliefs about others’ behaviours. Specifically, we show that inaccurate beliefs about other players’ behaviours can foster and stabilise cooperation via social norms. To show this, we combine a community's population dynamics with the behavioural dynamics of their members. In our model, individuals can join a community based on beliefs generated by public signals regarding the level of cooperation within, and decide to cooperate or not depending on these beliefs. These signals may overstate how much cooperation there really is. We show that even if individuals eventually learn the true level of cooperation, the initially false beliefs can trigger a dynamic that sustains high levels of cooperation. We also characterise how the rates of joining, leaving and learning in the community affect the cooperation level and community size simultaneously. Our results illustrate how false beliefs and social norms can help build cooperative communities.

The mate guarding theory of conservative clothing posits that veiling reduces women's physical allure and sexual attractiveness, thereby diminishing men's attraction towards them and deterring potential rivals for a woman's partner. This theory argues that the importance of veiling is influenced by ecological factors in a way that it is of higher importance to control women's sexuality in harsher environments to secure paternal investment. A prediction of this theory is that the importance of veiling should be influenced by community size, where individuals’ reputations, specifically men's, might have different weightings, and their perceived sense of controlling a partner's activity may differ. Using pre-existing data from seven countries encompassing over 9000 individuals, the current study explored the association of town size and importance of veiling for women. Results showed a U-shaped relationship where in small towns and large cities, individuals, specifically men, give more importance to the veiling of women. This finding not only has multiple implications in terms of the effect of community size on male policing behaviours of women and sexual restrictions, but it also might point to a wider relationship regarding the association of community size and moral values.

Major developments in the field of genetics in the past few decades have revolutionised notions of what it means to be human. Although currently only a few populations around the world practise a hunting and gathering lifestyle, this mode of subsistence has characterised members of our species since its very origins and allowed us to migrate across the planet. Therefore, the geographical distribution of hunter–gatherer populations, dependence on local ecosystems and connections to past populations and neighbouring groups have provided unique insights into our evolutionary origins. However, given the vulnerable status of hunter–gatherers worldwide, the development of the field of anthropological genetics requires that we reevaluate how we conduct research with these communities. Here, we review how the inclusion of hunter–gatherer populations in genetics studies has advanced our understanding of human origins, ancient population migrations and interactions as well as phenotypic adaptations and adaptability to different environments, and the important scientific and medical applications of these advancements. At the same time, we highlight the necessity to address yet unresolved questions and identify areas in which the field may benefit from improvements.

The prevalence of divorce in both parental and grandparental generations has led to a rise in the number of children who now have families that include both biological and step-grandparents. Despite the thorough examination of biological grandparents’ contributions in the recent literature, there remains a scarcity of studies focusing on the investment of step-grandparents. Using population-based data from a sample of 2494 parents in Germany, we assessed grandparental investment through financial support and assistance with childcare of grandparents (N = 4238) and step-grandparents (N = 486). The study revealed that step-grandparents provided lower levels of investment in their grandchildren compared with biological grandparents. Furthermore, the study identified that a longer duration of co-residence between step-grandparents and parents earlier in life did not correspond to an increase or decrease in step-grandparental investment. However, investment by separated biological grandparents increased with the increasing length of co-residence with parents. In line with the scarce literature on step-grandparental investment, these findings indicate that mating effort may be the most important motivation for step-grandparental investment.

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.

While humans are highly cooperative, they can also behave spitefully. Yet spite remains understudied. Spite can be normatively driven and while previous experiments have found some evidence that cooperation and punishment may spread via social learning, no experiments have considered the social transmission of spiteful behaviour. Here we present an online experiment where, following an opportunity to earn wealth, we asked participants to choose an action towards an anonymous partner across a full spectrum of social behaviour, from spite to altruism. In accordance with cultural evolutionary theory, participants were presented with social information that varied in source and content. Across six conditions, we informed participants that either the majority or the highest earner had chosen to behave spitefully, neutrally or altruistically. We found an overall tendency towards altruism, but at lower levels among those exposed to spite compared with altruism. We found no difference between social information that came from the majority or the highest earner. Exploratory analysis revealed that participants’ earnings negatively correlated with altruistic behaviour. Our results contrast with previous literature that report high rates of spite in experimental samples and a greater propensity for individuals to copy successful individuals over the majority.

Hunter–gatherer populations underwent a mass extinction in the Neolithic, and in present times face challenges such as explicit sedentarisation policies. An exception is in Nepal, where the nomadic Raute people receive monthly governmental individual payments. One consequence of the money transfers has been a significant increase in alcohol consumption, with nearly all individuals drinking industrially produced alcohol. Here we investigate the Raute demography based on a full census of 144 individuals. We show that the Raute exhibit the short life expectancies typical of other hunter–gatherer populations from Africa, Asia and America. Bayesian survival trajectory analysis demonstrated that heavy drinking by either parent substantially reduces offspring survival to age 15. Bayesian regressions revealed a significant effect of heavy drinking on maternal fertility by decreasing the number of living children and reducing the proportion of live children at the end of maternal reproductive life. Although the absence of data prior to monetary support precludes a direct assessment of long-term demographic trends, relatively stable population sizes over the last decades and a fertility rate close to the replacement rate rule out an imminent population crash. Further studies are required to elucidate the Rautes’ origins and relationship with other nomadic people in the region.

Loanword analysis is a unique contribution of historical linguistics to our understanding of prehistoric cultural interfaces. As language reflects the lives of its speakers, the substantiation of loanwords draws on the composite evidence from linguistic as well as auxiliary data from archaeology and genetics through triangulation. The Bronze Age of Central Asia is in principle linguistically mute, but a host of recent independent observations that tie languages, cultures and genetics together in various ways invites a comprehensive reassessment of six highly diagnostic loanwords (‘seven’, ‘name/fame’, ‘sister-in-law’, ‘honey’, ‘metal’ and ‘horse’) that are associated with the Bronze Age. Moreover, they are shared between Indo-European, Uralic, Turkic and sometimes Old Chinese. The successful identification of the interfaces for these loanwords can help settle longstanding debates on languages, migrations and the items themselves. Each item is analysed using the comparative method with reference to the archaeological record to assess the plausibility of a transfer. I argue that the six items can be dated to have entered Central and East Asian languages from immigrant Indo-European languages spoken in the Afanasievo and Andronovo cultures, including a novel source for the ‘horse’ in Old Chinese.

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.