The fossil record is scarce and incomplete by nature. Animals and ecological processes devour soft tissue and important bony details over time and, when the dust settles, we are faced with a patchy record full of variation. Fossil taxa are usually defined by craniodental characteristics, so unless postcranial bones are found associated with a skull, assignment to taxon is unstable. Naming a locomotor category based on fossil bone morphology by analogy to living hominoids is not uncommon, and when no single locomotor label fits, postcrania are often described as exhibiting a ‘mosaic’ of traits. Here, we contend that the unavoidable variation that characterises the fossil record can be described far more rigorously based on extensive work in human neurobiology and neuroanatomy, movement sciences and motor control and biomechanics research. In neurobiology, degeneracy is a natural mechanism of adaptation allowing system elements that are structurally different to perform the same function. This concept differs from redundancy as understood in engineering, where the same function is performed by identical elements. Assuming degeneracy, structurally different elements are able to produce different outputs in a range of environmental contexts, favouring ecological robusticity by enabling adaptations. Furthermore, as degeneracy extends to genome level, genetic variation is sustained, so that genes which might benefit an organism in a different environment remain part of the genome, favouring species’ evolvability.

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.

Hunter–gatherer occupations of small islands are rare in world prehistory and it is widely accepted that island settlement is facilitated by agriculture. The Ryukyu Islands contradict that understanding on two counts: not only did they have a long history of hunter–gatherer settlement, but they also have a very late date for the onset of agriculture, which only reached the archipelago between the eighth and thirteenth centuries AD. Here, we combine archaeology and linguistics to propose a tripartite model for the spread of agriculture and Ryukyuan languages to the Ryukyu Islands. Employing demographic growth, trade/piracy and the political influence of neighbouring states, this model provides a synthetic yet flexible understanding of farming/language dispersals in the Ryukyus within the complex historical background of medieval East Asia.

Associative Tool Use (ATU) describes the use of two or more tools in combination, with the literature further differentiating between Tool set use, Tool composite use, Sequential tool use and Secondary tool use. Research investigating the cognitive processes underlying ATU has shown that some primate and bird species spontaneously invent Tool set and Sequential tool use. Yet studies with humans are sparse. Whether children are also able to spontaneously invent ATU behaviours and at what age this ability emerges is poorly understood. We addressed this gap in the literature with two experiments involving preschoolers (E1, N = 66, 3 years 6 months to 4 years 9 months; E2, N = 119, 3 years 0 months to 6 years 10 months) who were administered novel tasks measuring Tool set, Metatool and Sequential tool use. Participants needed to solve the tasks individually, without the opportunity for social learning (except for enhancement effects). Children from 3 years of age spontaneously invented all of the types of investigated ATU behaviours. Success rates were low, suggesting that individual invention of ATU in novel tasks is still challenging for preschoolers. We discuss how future studies can use and expand our tasks to deepen our understanding of tool use and problem-solving in humans and non-human animals.

Musculoskeletal models, like all theoretical models of physical processes, depend on the assumptions needed to construct the model. For musculoskeletal models, these assumptions include, among other things, the kinematic data, the kinetic data and the muscle parameters. The former (dynamic) data can be acquired relatively easily from living subjects, but the latter are usually based on limited information, frequently determined from cadaver studies performed on elderly individuals. Previously, we determined the sensitivity of forces to dynamic differences among 10 humans walking on a straight path. Here, we assess the sensitivity of the muscle and joint reaction forces developed in human walking to variable muscle parameters obtained from 10 living adults, whose data were recently reported, and compared the results with the values from a standard model that depends on cadaveric data. We found that, while the force patterns across the stance cycle were similar among muscle parameter models, differences of as much as 15% in the force magnitude were produced. Whether or not the variation between the standard model and other muscle parameters is important depends on why the forces are required.

Evolutionary demographers often invoke tradeoffs between reproduction and survival to explain reductions in fertility during demographic transitions. The evidence for such tradeoffs in humans has been mixed, partly because tradeoffs may be masked by individual differences in quality or access to resources. Unmasking tradeoffs despite such phenotypic correlations requires sophisticated statistical analyses that account for endogeneity among variables and individual differences in access to resources. Here we tested for costs of reproduction using N = 13,663 birth records from the maternity hospital in Basel, Switzerland, 1896–1939, a period characterised by rapid fertility declines. We predicted that higher parity is associated with worse maternal and offspring condition at the time of birth, adjusting for age and a variety of covariates. We used Bayesian multivariate, multilevel models to simultaneously analyse multiple related outcomes while accounting for endogeneity, appropriately modelling non-linear effects, dealing with hierarchical data structures, and effectively imputing missing data. Despite all these efforts, we found virtually no evidence for costs of reproduction. Instead, women with better access to resources had fewer children. Barring limitations of the data, these results are consistent with demographic transitions reflecting women's investment in their own embodied capital and/or the adoption of maladaptive low-fertility norms by elites.

Theories of early cooperation in human society often draw from a small sample of ethnographic studies of surviving populations of hunter–gatherers, most of which are now sedentary. Borneo hunter–gatherers (Punan, Penan) have seldom figured in comparative research because of a decades-old controversy about whether they are the descendants of farmers who adopted a hunting and gathering way of life. In 2018 we began an ethnographic study of a group of still-nomadic hunter–gatherers who call themselves Punan Batu (Cave Punan). Our genetic analysis clearly indicates that they are very unlikely to be the descendants of neighbouring agriculturalists. They also preserve a song language that is unrelated to other languages of Borneo. Dispersed travelling groups of Punan Batu with fluid membership use message sticks to stay in contact, co-operate and share resources as they journey between rock shelters and forest camps. Message sticks were once widespread among nomadic Punan in Borneo, but have largely disappeared in sedentary Punan villages. Thus the small community of Punan Batu offers a rare glimpse of a hunting and gathering way of life that was once widespread in the forests of Borneo, where prosocial behaviour extended beyond the face-to-face community, facilitating successful collective adaptation to the diverse resources of Borneo's forests.

Why do we leak lubricant from the eyes to solicit comfort from others? Why do we bare our teeth and crinkle our faces to express non-aggression? The defensive mimic theory proposes that a broad range of human emotional expressions evolved originally as exaggerated, temporally extended mimics of the fast, defensive reflexes that normally protect the body surface. Defensive reflexes are so important to survival that they cannot be safely suppressed; yet they also broadcast information about an animal's internal state, information that can potentially be exploited by other animals. Once others can observe and exploit an animal's defensive reflexes, it may be advantageous to the animal to run interference by creating mimic defensive actions, thereby manipulating the behaviour of others. Through this interaction over millions of years, many human emotional expressions may have evolved. Here, human social signals including smiling, laughing and crying, are compared component-by-component with the known, well-studied features of primate defensive reflexes. It is suggested that the defensive mimic theory can adequately account for the physical form of not all, but a large range of, human emotional expression.

This paper surveys five human societal types – mobile foragers, horticulturalists, pre-state agriculturalists, state-based agriculturalists and liberal democracies – from the perspective of three core social problems faced by interacting individuals: coordination problems, social dilemmas and contest problems. We characterise the occurrence of these problems in the different societal types and enquire into the main force keeping societies together given the prevalence of these. To address this, we consider the social problems in light of the theory of repeated games, and delineate the role of intertemporal incentives in sustaining cooperative behaviour through the reciprocity principle. We analyse the population, economic and political structural features of the five societal types, and show that intertemporal incentives have been adapted to the changes in scope and scale of the core social problems as societies have grown in size. In all societies, reciprocity mechanisms appear to solve the social problems by enabling lifetime direct benefits to individuals for cooperation. Our analysis leads us to predict that as societies increase in complexity, they need more of the following four features to enable the scalability and adaptability of the reciprocity principle: nested grouping, decentralised enforcement and local information, centralised enforcement and coercive power, and formal rules.

Bipedal locomotion is a hallmark of being human. Yet the body form from which bipedalism evolved remains unclear. Specifically, the positional behaviour (i.e. orthograde vs. pronograde) and the length of the lumbar spine (i.e. long and mobile vs. short and stiff) of the last common ancestor (LCA) of the African great apes and humans require further investigation. While fossil evidence would be the most conclusive, the paucity of hominid fossils from 5–10 million years ago makes this field of research challenging. In their absence, extant primate anatomy and behaviour may offer some insight into the ancestral body form from which bipedalism could most easily evolve. Here, we quantify the frequency of bipedalism in a large sample (N = 496) of zoo-housed hominoids and cercopithecines. Our results show that while each studied species of ape and monkey can move bipedally, hylobatids are significantly more bipedal and engage in bipedal locomotion more frequently and for greater distances than any other primate sampled. These data support hypotheses of an orthograde, long-backed and arboreal LCA, which is consistent with hominoid fossils from the middle-to-late Miocene. If true, knuckle-walking evolved in parallel in Pan and Gorilla, and the human body form, particularly the long lower back and orthograde posture, is conserved.

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.

Research among non-industrial societies suggests that body kinematics adopted during running vary between groups according to the cultural importance of running. Among groups in which running is common and an important part of cultural identity, runners tend to adopt what exercise scientists and coaches consider to be good technique for avoiding injury and maximising performance. In contrast, among groups in which running is not particularly culturally important, people tend to adopt suboptimal technique. This paper begins by describing key elements of good running technique, including landing with a forefoot or midfoot strike pattern and leg oriented roughly vertically. Next, we review evidence from non-industrial societies that cultural attitudes about running associate with variation in running techniques. Then, we present new data from Tsimane forager–horticulturalists in Bolivia. Our findings suggest that running is neither a common activity among the Tsimane nor is it considered an important part of cultural identity. We also demonstrate that when Tsimane do run, they tend to use suboptimal technique, specifically landing with a rearfoot strike pattern and leg protracted ahead of the knee (called overstriding). Finally, we discuss processes by which culture might influence variation in running techniques among non-industrial societies, including self-optimisation and social learning.

Research on paternal investment and child growth and development is limited outside of high-income countries. Using nationally representative data from low-resource Serbian Roma communities, this study examined father investment (direct care), its predictors and the associations between paternal investment, stepfather presence and child physical growth and early development. The sample included 1222 children aged 35–59 months, out of which 235 were living with biological fathers. Child outcomes included height-for-age Z-scores, stunting and early child developmental score. Roma paternal investment was relatively low. There was a positive association of father investment and children's height, and no association with developmental score. The presence of father vs. stepfather did not exert any influence on children. Instead, maternal and child characteristics explained both the overall development and height for Roma children. Thus, older children, born to literate, lower parity mothers of higher status and greater investment had better developmental and growth outcomes; girls were the preferred sex, owing to expected fitness benefits. Reverse causality emerged as the most likely pathway through which the cross-sectional association of father direct care with child growth may manifest, such that Roma fathers tend to bias their investment towards taller, more endowed children, because of greater fitness pay-off.

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.

Professional religious specialists centralised religious authority in early human societies and represented some of the earliest instances of formalised social leadership. These individuals played a central role in the emergence of organised religion and transitions to more stratified human societies. Evolutionary theories highlight a range of environmental, economic and social factors that are potentially causally related to the emergence of professional religious specialists in human history. There remains little consensus over the relative importance of these factors and whether professional religious specialists were the outcome or driver of increased socio-cultural complexity. We built a global dataset of hunter–gatherer societies and developed a novel method of exploratory phylogenetic path analysis. This enabled us to systematically identify the factors associated with the emergence of professional religious specialists and infer the directionality of causal dependencies. We find that environmental predictability, environmental richness, pathogen load, social leadership and food storage systems are all correlated with professional religious specialists. However, only food storage is directly related to the emergence of professional religious specialists. Our findings are most consistent with the claim that the early stages of organised religion were the outcome rather than driver of increased socio-economic complexity.

High-dimensional datasets on cultural characters contribute to uncovering insights about factors that influence cultural evolution. Because cultural variation in part reflects descent processes with a hierarchical structure – including the descent of populations and vertical transmission of cultural traits – methods designed for hierarchically structured data have potential to find applications in the analysis of cultural variation. We adapt a network-based hierarchical clustering method for use in analysing cultural variation. Given a set of entities, the method constructs a similarity network, hierarchically depicting community structure among them. We illustrate the approach using four datasets: pronunciation variation in the US mid-Atlantic region, folklore variation in worldwide cultures, phonemic variation across worldwide languages and temporal variation in first names in the US. In these examples, the method provides insights into processes that affect cultural variation, uncovering geographic and other influences on observed patterns and cultural characters that make important contributions to them.

Laboratory-based studies indicate that a major evolutionary advantage of bipedalism is enabling humans to walk with relatively low energy expenditure. However, such studies typically record subjects walking on even surfaces or treadmills that do not represent the irregular terrains our species encounters in natural environments. To date, few studies have quantified walking kinematics on natural terrains. Here we used high-speed video to record marker-based kinematics of 21 individuals from a Tsimane forager–horticulturalist community in the Bolivian Amazon walking on three different terrains: a dirt field, a forest trail and an unbroken forest transect. Compared with the field, in the unbroken forest participants contacted the ground with more protracted legs and flatter foot postures, had more inclined trunks, more flexed hips and knees, and raised their feet higher during leg swing. In contrast, kinematics were generally similar between trail and field walking. These results provide preliminary support for the idea that irregular natural surfaces like those in forests cause humans to alter their walking kinematics, such that travel in these environments could be more energetically expensive than would be assumed from laboratory-based data. These findings have important implications for the evolutionary energetics of human foraging in environments with challenging terrains.

Credibility Enhancing Displays have been shown to be an important component in the transmission of empirically unverifiable cultural content such as religious beliefs. Decreased Credibility Enhancing Displays are a major predictor of religious decline. However, because declines in belief are often paired with the decreasing importance of religious institutions, existing research has not yet shown the effect of Credibility Enhancing Displays as separate from this institutional decline. Here, we assess the role of past Credibility Enhancing Display exposure among the baptised Catholic population of Ireland in predicting who retains a Catholic identity and religious beliefs among those who reject the Catholic Church. We find that leaving Catholicism outright (i.e. ‘ex-Catholicism’) is predicted by low Credibility Enhancing Display exposure, but rejecting the Church while retaining a Catholic identity (i.e. ‘liminal Catholicism’) and theistic belief is not. High perceived prevalence of clerical paedophiles (i.e. religious hypocrisy) predicts both groups similarly. Higher exposure to Credibility Enhancing Displays predicts higher orthodox Catholic beliefs and Catholic morality among Catholics, but with inconsistent and even negative effects among the other groups. High perceived prevalence of clerical paedophiles predicts the rejection of orthodox Catholic beliefs, but not the rejection of theism or a Catholic identity.