Using machine learning on new feature sets extracted from three-dimensional models of broken animal bones to classify fragments according to break agent.

Distinguishing agents of bone modification at paleoanthropological sites is an important means of understanding early hominin evolution. Fracture pattern analysis is used to help determine site formation processes, including whether hominins were hunting or scavenging for animal food resources. Determination of how these behaviors manifested in ancient human sites has major implications for our biological and behavioral evolution, including social and cognitive abilities, dietary impacts of having access to in-bone nutrients like marrow, and cultural variation in butchering and food processing practices. Nevertheless, previous analyses remain inconclusive, often suffering from lack of replicability, misuse of mathematical methods, and/or failure to overcome equifinality. In this paper, we present a new approach aimed at distinguishing bone fragments resulting from hominin and carnivore breakage. Our analysis is founded on a large collection of scanned three-dimensional models of fragmentary bone broken by known agents, to which we apply state of the art machine learning algorithms. Our classification of fragments achieves an average mean accuracy of 77% across tests, thus demonstrating notable, but not overwhelming, success for distinguishing the agent of breakage. We note that, while previous research applying such algorithms has claimed higher success rates, fundamental errors in the application of machine learning protocols suggest that the reported accuracies are unjustified and unreliable. The systematic, fully documented, and proper application of machine learning algorithms leads to an inherent reproducibility of our study, and therefore our methods hold great potential for deciphering when and where hominins first began exploiting marrow and meat, and clarifying their importance and influence on human evolution.

Modeling Oldowan tool transport from a primate perspective.

Living nonhuman primates have long served as a referential framework for understanding various aspects of hominin biological and cultural evolution. Comparing the cognitive, social, and ecological contexts of nonhuman primate and hominin tool use has allowed researchers to identify key adaptations relevant to the evolution of hominin behavior. Although the Oldowan is often considered to be a major evolutionary milestone, it has been argued that the Oldowan is rather an extension of behaviors already present in the ape lineage. This is based on the fact that while apes move tools through repeated, unplanned, short-distance transport bouts, they produce material patterning often associated with long-distance transport, planning, and foresight in the Oldowan. Nevertheless, remain fundamental differences in how Oldowan core and flake technology and nonhuman primate tools are used. The goal of the Oldowan hominins is to produce sharp-edged flakes, whereas nonhuman primates use stone tools primarily as percussors. Here, we present an agent-based model that investigates the explanatory power of the ape tool transport model in light of these differences. The model simulates the formation of the Oldowan record under the conditions of an accumulated short-distance transport pattern, as seen in extant chimpanzees. Our results show that while ape tool transport can account for some of the variation observed in the archaeological record, factors related to use-life duration severely limit how far an Oldowan core can be moved through repeated short-distance transport bouts. Thus, the ape tool transport has limitations in its ability to explain patterns in the Oldowan. These results provide a basis for discussing adaptive processes that would have facilitated the development of the Oldowan.

Social cognition and metacognition in great apes: a theory.

Twenty-five years ago, at the founding of this journal, there existed only a few conflicting findings about great apes' social-cognitive skills (theory of mind). In the 2 ½ decades since, we have discovered that great apes understand the goals, intentions, perceptions, and knowledge of others, and they use this knowledge to their advantage in competitive interactions. Twenty-five years ago there existed basically no studies on great apes' metacognitive skills. In the 2 ½ decades since, we have discovered that great apes monitor their uncertainty and base their decisions on that, or else decide to gather more information to make better decisions. The current paper reviews the past 25 years of research on great ape social cognition and metacognition and proposes a theory about how the two are evolutionarily related.

Signatures of adaptive evolution in platyrrhine primate genomes.

The platyrrhine family Cebidae (capuchin and squirrel monkeys) exhibit among the largest primate encephalization quotients. Each cebid lineage is also characterized by notable lineage-specific traits, with capuchins showing striking similarities to Hominidae such as high sensorimotor intelligence with tool use, advanced cognitive abilities, and behavioral flexibility. Here, we take a comparative genomics approach, performing genome-wide tests for positive selection across five cebid branches, to gain insight into major periods of cebid adaptive evolution. We uncover candidate targets of selection across cebid evolutionary history that may underlie the emergence of lineage-specific traits. Our analyses highlight shifting and sustained selective pressures on genes related to brain development, longevity, reproduction, and morphology, including evidence for cumulative and diversifying neurobiological adaptations across cebid evolution. In addition to generating a high-quality reference genome assembly for robust capuchins, our results lend to a better understanding of the adaptive diversification of this distinctive primate clade.

Hierarchical object combination and tool use in the great apes and human children.

Object manipulation can be used as a comparative scale of cognitive development among primates, including humans. Combinatory object manipulation is a precursor of tool-using behavior that indicates material intelligence in primates. However, developmental data on it regarding the great apes other than chimpanzees is insufficient. We conducted a longitudinal investigation of humans and chimpanzees as well as a cross-sectional examination of other great-ape infants (two bonobos, three gorillas, and four orangutans) in captive settings by using two kinds of tasks that required either inserting or stacking combinatory action. The four species of great apes and humans demonstrated both types of combinatory object manipulation during infancy. However, the order of development in different types of combinatory object manipulations varied among the great apes. Furthermore, we applied a nesting-cup task to examine the hierarchical complexity in the combinatory strategies of human children and adult chimpanzees. Both of them exhibited highly hierarchical combinations in the nesting-cup task and employed the subassembly strategy, indicating that an action merge may exist not only in human children but also in adult chimpanzees. The results were discussed with reviews of the tool-use literature from the wild great apes. The early acquisition of an inserting action in the chimpanzees may explain the tool utilization commonality reported in wild chimpanzees. The combinatory object manipulation may have worked as an external enhancer to achieve an additional hierarchical complexity in cognition and behavior, eventually leading humans to develop a language system.

Learning from communication versus observation in great apes.

When human infants are intentionally addressed by others, they tend to interpret the information communicated as being relevant to them and worth acquiring. For humans, this attribution of relevance leads to a preference to learn from communication, making it possible to accumulate knowledge over generations. Great apes are sensitive to communicative cues, but do these cues also activate an expectation of relevance? In an observational learning paradigm, we demonstrated to a sample of nonhuman great apes (bonobos, chimpanzees, orangutans; N = 24) how to operate on a food dispenser device. When apes had the opportunity to choose between an effective and an ineffective method in the baseline conditions, the majority of them chose the effective method. However, when the ineffective method was demonstrated in a communicative way, they failed to prioritize efficiency, even though they were equally attentive in both conditions. This suggests that the ostensive demonstration elicited an expectation of relevance that modified apes' interpretation of the situation, potentially leading to a preference to learn from communication, as human children do.

Anthropological Prosociality via Sub-Group Level Selection.

A perennial challenge of evolutionary psychology is explaining prosocial traits such as a preference for fairness rather than inequality, compassion towards suffering, and an instinctive ability to coordinate within small teams. Considering recent fossil evidence and a novel logical test, we deem present explanations insufficiently explanatory of the divergence of hominins. In answering this question, we focus on the divergence of hominins from the last common ancestor (LCA) shared with Pan. We consider recent fossil discoveries that indicate the LCA was bipedal, which reduces the cogency of this explanation for hominin development. We also review evolutionary theory that claims to explain how hominins developed into modern humans, however it is found that no mechanism differentiates hominins from other primates. Either the mechanism was available to the last common ancestor (LCA) (with P. troglodytes as its proxy), or because early hominins had insufficient cognition to utilise the mechanism. A novel mechanism, sub-group level selection (sGLS) is hypothesised by triangulating two pieces of data rarely considered by evolutionary biologists. These are behavioural dimorphism of Pan (chimpanzees and bonobos) that remain identifiable in modern humans, and the social behaviour of primate troops in a savannah ecology. We then contend that sGLS supplied an exponential effect which was available to LCA who left the forest, but was not sufficiently available to any other primates. In conclusion, while only indirectly supported by various evidence, sGLS is found to be singularly and persuasively explanatory of human's unique evolutionary story.

Social pressure drives "conversational rules" in great apes.

In the last decade, two hypotheses, one on the evolution of animal vocal communication in general and the other on the origins of human language, have gained ground. The first hypothesis argues that the complexity of communication co-evolved with the complexity of sociality. Species forming larger groups with complex social networks have more elaborate vocal repertoires. The second hypothesis posits that the core of communication is represented not only by what can be expressed by an isolated caller, but also by the way that vocal interactions are structured, language being above all a social act. Primitive forms of conversational rules based on a vocal turn-taking principle are thought to exist in primates. To support and bring together these hypotheses, more comparative studies of socially diverse species at different levels of the primate phylogeny are needed. However, the majority of available studies focus on monkeys, primates that are distant from the human lineage. Great apes represent excellent candidates for such comparative studies because of their phylogenetic proximity to humans and their varied social lives. We propose that studying vocal turn-taking in apes could address several major gaps regarding the social relevance of vocal turn-taking and the evolutionary trajectory of this behaviour among anthropoids. Indeed, how the social structure of a species may influence the vocal interaction patterns observed among group members remains an open question. We gathered data from the literature as well as original unpublished data (where absent in the literature) on four great ape species: chimpanzees Pan troglodytes, bonobos Pan paniscus, western lowland gorillas Gorilla gorilla gorilla and Bornean orang-utans Pongo pygmaeus. We found no clear-cut relationship between classical social complexity metrics (e.g. number of group members, interaction rates) and vocal complexity parameters (e.g. repertoire size, call rates). Nevertheless, the nature of the society (i.e. group composition, diversity and valence of social bonds) and the type of vocal interaction patterns (isolated calling, call overlap, turn-taking-based vocal exchanges) do appear to be related. Isolated calling is the main vocal pattern found in the species with the smallest social networks (orang-utan), while the other species show vocal interactions that are structured according to temporal rules. A high proportion of overlapping vocalisations is found in the most competitive species (chimpanzee), while vocal turn-taking predominates in more tolerant bonobos and gorillas. Also, preferentially interacting individuals and call types used to interact are not randomly distributed. Vocal overlap ('chorusing') and vocal exchange ('conversing') appear as possible social strategies used to advertise/strengthen social bonds. Our analyses highlight that: (i) vocal turn-taking is also observed in non-human great apes, revealing universal rules for conversing that may be deeply rooted in the primate lineage; (ii) vocal interaction patterns match the species' social lifestyle; (iii) although limited to four species here, adopting a targeted comparative approach could help to identify the multiple and subtle factors underlying social and vocal complexity. We believe that vocal interaction patterns form the basis of a promising field of investigation that may ultimately improve our understanding of the socially driven evolution of communication.

Emergence of perceptuomotor relationships during paleolithic stone toolmaking learning: intersections of observation and practice.

Stone toolmaking is a human motor skill which provides the earliest archeological evidence motor skill and social learning. Intentionally shaping a stone into a functional tool relies on the interaction of action observation and practice to support motor skill acquisition. The emergence of adaptive and efficient visuomotor processes during motor learning of such a novel motor skill requiring complex semantic understanding, like stone toolmaking, is not understood. Through the examination of eye movements and motor skill, the current study sought to evaluate the changes and relationship in perceptuomotor processes during motor learning and performance over 90 h of training. Participants' gaze and motor performance were assessed before, during and following training. Gaze patterns reveal a transition from initially high gaze variability during initial observation to lower gaze variability after training. Perceptual changes were strongly associated with motor performance improvements suggesting a coupling of perceptual and motor processes during motor learning.

The early hunting dog from Dmanisi with comments on the social behaviour in Canidae and hominins.

The renowned site of Dmanisi in Georgia, southern Caucasus (ca. 1.8 Ma) yielded the earliest direct evidence of hominin presence out of Africa. In this paper, we report on the first record of a large-sized canid from this site, namely dentognathic remains, referable to a young adult individual that displays hypercarnivorous features (e.g., the reduction of the m1 metaconid and entoconid) that allow us to include these specimens in the hypodigm of the late Early Pleistocene species Canis (Xenocyon) lycaonoides. Much fossil evidence suggests that this species was a cooperative pack-hunter that, unlike other large-sized canids, was capable of social care toward kin and non-kin members of its group. This rather derived hypercarnivorous canid, which has an East Asian origin, shows one of its earliest records at Dmanisi in the Caucasus, at the gates of Europe. Interestingly, its dispersal from Asia to Europe and Africa followed a parallel route to that of hominins, but in the opposite direction. Hominins and hunting dogs, both recorded in Dmanisi at the beginning of their dispersal across the Old World, are the only two Early Pleistocene mammal species with proved altruistic behaviour towards their group members, an issue discussed over more than one century in evolutionary biology.

No evidence for widespread island extinctions after Pleistocene hominin arrival.

The arrival of modern humans into previously unoccupied island ecosystems is closely linked to widespread extinction, and a key reason cited for Pleistocene megafauna extinction is anthropogenic overhunting. A common assumption based on late Holocene records is that humans always negatively impact insular biotas, which requires an extrapolation of recent human behavior and technology into the archaeological past. Hominins have been on islands since at least the early Pleistocene and Homo sapiens for at least 50 thousand y (ka). Over such lengthy intervals it is scarcely surprising that significant evolutionary, behavioral, and cultural changes occurred. However, the deep-time link between human arrival and island extinctions has never been explored globally. Here, we examine archaeological and paleontological records of all Pleistocene islands with a documented hominin presence to examine whether humans have always been destructive agents. We show that extinctions at a global level cannot be associated with Pleistocene hominin arrival based on current data and are difficult to disentangle from records of environmental change. It is not until the Holocene that large-scale changes in technology, dispersal, demography, and human behavior visibly affect island ecosystems. The extinction acceleration we are currently experiencing is thus not inherent but rather part of a more recent cultural complex.

Comparing prehistoric constructed languages: world-building and its role in understanding prehistoric languages.

In this paper, we compare the languages each of the authors invented as prehistoric languages for popular culture media. Schreyer's language, Beama, was created for the film Alpha (2018), while Adger's language, Tan!aa Kawawa ki, was created for a television series on how early hominins spread throughout the world (the series was green-lit but then cancelled). We argue that though this creative process may seem far removed from classical research paradigms on language evolution, it can provide some insight into how disparate research on the possible properties of prehistoric languages can be brought together to illustrate how these languages might have worked as whole linguistic systems within these imagined worlds, as well as in prehistory. This article is part of the theme issue 'Reconstructing prehistoric languages'.

Persuasive conversation as a new form of communication in Homo sapiens.

The aim of this paper is twofold: to propose that conversation is the distinctive feature of Homo sapiens' communication; and to show that the emergence of modern language is tied to the transition from pantomime to verbal and grammatically complex forms of narrative. It is suggested that (animal and human) communication is a form of persuasion and that storytelling was the best tool developed by humans to convince others. In the early stage of communication, archaic hominins used forms of pantomimic storytelling to persuade others. Although pantomime is a powerful tool for persuasive communication, it is proposed that it is not an effective tool for persuasive conversation: conversation is characterized by a form of reciprocal persuasion among peers; instead, pantomime has a mainly asymmetrical character. The selective pressure towards persuasive reciprocity of the conversational level is the evolutionary reason that allowed the transition from pantomime to grammatically complex codes in H. sapiens, which favoured the evolution of speech. This article is part of the theme issue 'Reconstructing prehistoric languages'.

Non-human primates use combined rules when deciding under ambiguity.

Decision outcomes in unpredictable environments may not have exact known probabilities. Yet the predictability level of outcomes matters in decisions, and animals, including humans, generally avoid ambiguous options. Managing ambiguity may be more challenging and requires stronger cognitive skills than decision-making under risk, where decisions involve known probabilities. Here we compare decision-making in capuchins, macaques, orangutans, gorillas, chimpanzees and bonobos in risky and ambiguous contexts. Subjects were shown lotteries (a tray of potential rewards, some large, some small) and could gamble a medium-sized food item to obtain one of the displayed rewards. The odds of winning and losing varied and were accessible in the risky context (all rewards were visible) or partially available in the ambiguous context (some rewards were covered). In the latter case, the level of information varied from fully ambiguous (individuals could not guess what was under the covers) to predictable (individuals could guess). None of the species avoided gambling in ambiguous lotteries and gambling rates were high if at least two large rewards were visible. Capuchins and bonobos ignored the covered items and gorillas and macaques took the presence of potential rewards into account, but only chimpanzees and orangutans could consistently build correct expectations about the size of the covered rewards. Chimpanzees and orangutans combined decision rules according to the number of large visible rewards and the level of predictability, a process resembling conditional probabilities assessment in humans. Despite a low sample size, this is the first evidence in non-human primates that a combination of several rules can underlie choices made in an unpredictable environment. Our finding that non-human primates can deal with the uncertainty of an outcome when exchanging one food item for another is a key element to the understanding of the evolutionary origins of economic behaviour. This article is part of the theme issue 'Existence and prevalence of economic behaviours among non-human primates'.

Giraffes and hominins: reductionist model predictions of compressive loads at the spine base for erect exponents of the animal kingdom.

In humans, compressive stress on intervertebral discs is commonly deployed as a measurand for assessing the loads that act within the spine. Examining this physical quantity is crucially beneficial: the intradiscal pressure can be directly measured in vivo in humans, and is immediately related to compressive stress. Hence, measured intradiscal pressure data are very useful for validating such biomechanical animal models that have the spine incorporated, and can, thus, compute compressive stress values. Here, we use human intradiscal pressure data to verify the predictions of a reductionist spine model, which has in fact only one joint degree of freedom. We calculate the pulling force of one lumped anatomical structure that acts past this (intervertebral) joint at the base of the spine, lumbar in hominins, cervical in giraffes, to compensate the torque that is induced by the weight of all masses located cranially to the base. Given morphometric estimates of the human and australopith trunks, respectively, and the giraffe's neck, as well as the respective structures' lever arms and disc areas, we predict, for all three species, the compressive stress on the intervertebral disc at the spine base, while systematically varying the angular orientation of the species' spinal columns with respect to gravity. The comparison between these species demonstrates that hominin everyday compressive disc stresses are lower than those in big quadrupedal animals. Within each species, erecting the spine from being bent forward by, for example, thirty degrees to fully upright posture reduces the compressive disc stress roughly to a third. We conclude that erecting the spine immediately allows the carrying of extra loads of the order of body weight, and yet the compressive disc stress is lower than in a moderately forward-bent posture with no extra load.

Human children, but not great apes, become socially closer by sharing an experience in common ground.

To create social closeness, humans engage in a variety of social activities centered around shared experiences. Even simply watching the same video side by side creates social closeness in adults and children. However, perhaps surprisingly, a similar psychological mechanism was recently shown in great apes. Here we asked whether the process by which this social closeness is created is the same for children and great apes. Each participant entered a room to see an experimenter (E1) watching a video. In one condition, E1 looked to the participant at the start of the video to establish common ground that they were watching the video together. In another condition, E1 did not look to the participant in this way so that the participant knew they were watching the same video, but the participant did not know whether E1 was aware of this as well, so there was no common ground (E1 looked to the participant later in the procedure). Children, but not great apes, approached the experimenter faster after the common ground condition, suggesting that although both humans and great apes create social closeness by co-attending to something in close proximity, creating social closeness by sharing experiences in common ground may be a uniquely human social-cognitive process.

Innovation, life history and social networks in human evolution.

There is a famous puzzle about the first 3 million years of archaeologically visible human technological history. The pace of change, of innovation and its uptake, is extraordinarily slow. In particular, the famous handaxes of the Acheulian technological tradition first appeared about 1.7 Ma, and persisted with little change until about 800 ka, perhaps even longer. In this paper, I will offer an explanation of that stasis based in the life history and network characteristics that we infer (on phylogenetic grounds) to have characterized earlier human species. The core ideas are that (i) especially in earlier periods of hominin evolution, we are likely to find archaeological traces only of widespread and persisting technologies and practices; (ii) the record is not a record of the rate of innovation, but the rate of innovations establishing in a landscape; (iii) innovations are extremely vulnerable to stochastic loss while confined to the communities in which they are made and established; (iv) the export of innovation from the local group is sharply constrained if there is a general pattern of hostility and suspicion between groups, or even if there is just little contact between adults of adjoining groups. That pattern is typical of great apes and likely, therefore, to have characterized at least early hominin social lives. Innovations are unlikely to spread by adult-to-adult interactions across community boundaries. (v) Chimpanzees and bonobos are characterized by male philopatry and subadult female dispersal; that is, therefore, the most likely early hominin pattern. If so, the only innovations at all likely to expand beyond the point of origin are those acquired by subadult females, and ones that can be expressed by those females, at high enough frequency and salience for them to spread, in the bands that the females join. These are very serious filters on the spread of innovation. This article is part of the theme issue 'Life history and learning: how childhood, caregiving and old age shape cognition and culture in humans and other animals'.

The emergence of emotionally modern humans: implications for language and learning.

According to the Cooperative Breeding Hypothesis, apes with the life-history attributes of those in the line leading to the genus Homo could not have evolved unless male and female allomothers had begun to help mothers care for and provision offspring. As proposed elsewhere, the unusual way hominins reared their young generated novel phenotypes subsequently subjected to Darwinian social selection favouring those young apes best at monitoring the intentions, mental states and preferences of others and most motivated to attract and appeal to caretakers. Not only were youngsters acquiring information in social contexts different from those of other apes, but they would also have been emotionally and neurophysiologically different from them in ways that are relevant to how humans learn. Contingently delivered rewards to dependents who attracted and ingratiated themselves with allomothers shaped their behaviours and vocalizations and transformed the way developing youngsters learned from others and internalized their preferences. This article is part of the theme issue 'Life history and learning: how childhood, caregiving and old age shape cognition and culture in humans and other animals'.

The human life history is adapted to exploit the adaptive advantages of culture.

Humans evolved from an ape ancestor that was highly intelligent, moderately social and moderately dependent on cultural adaptations for subsistence technology (tools). By the late Pleistocene, humans had become highly dependent on culture for subsistence and for rules to organize a complex social life. Adaptation by cultural traditions transformed our life history, leading to an extended juvenile period to learn subsistence and social skills, post-reproductive survival to help conserve and transmit skills, a dependence on social support for mothers of large-brained, very dependent and nutrient-demanding offspring, males devoting substantial effort to provisioning rather than mating, and the cultivation of large social networks to tap pools in information unavailable to less social species. One measure of the success of the exploitation of culture is that the minimum inter-birth interval of humans is nearly half that of our ape relatives. Another measure is the wide geographical distribution of humans compared with other apes, based on subsistence systems adapted to fine-scale spatial environmental variation. An important macro-evolutionary question is why our big-brained, culture-intensive life-history strategy evolved so recently and in only our lineage. We suggest that increasing spatial and temporal variation in the Pleistocene favoured cultural adaptations. This article is part of the theme issue 'Life history and learning: how childhood, caregiving and old age shape cognition and culture in humans and other animals'.