Phylogenetic differences in the morphology and shape of the central sulcus in great apes and humans: implications for the evolution of motor functions.

The central sulcus divides the primary motor and somatosensory cortices in many anthropoid primate brains. Differences exist in the surface area and depth of the central sulcus along the dorso-ventral plane in great apes and humans compared to other primate species. Within hominid species, there are variations in the depth and aspect of their hand motor area, or knob, within the precentral gyrus. In this study, we used post-image analyses on magnetic resonance images to characterize the central sulcus shape of humans, chimpanzees (Pan troglodytes), gorillas (Gorilla gorilla), and orangutans (Pongo pygmaeus and Pongo abelii). Using these data, we examined the morphological variability of central sulcus in hominids, focusing on the hand region, a significant change in human evolution. We show that the central sulcus shape differs between great ape species, but all show similar variations in the location of their hand knob. However, the prevalence of the knob location along the dorso-ventral plane and lateralization differs between species and the presence of a second ventral motor knob seems to be unique to humans. Humans and orangutans exhibit the most similar and complex central sulcus shapes. However, their similarities may reflect divergent evolutionary processes related to selection for different positional and habitual locomotor functions.

Hominin turnover at Laetoli is associated with vegetation change: Multiproxy evidence from the large herbivore community.

Vegetation change in eastern Africa during the Pliocene would have had an important impact on hominin adaptation and ecology, and it may have been a key driver of hominin macroevolution, including the extinction of Australopithecus and the emergence of Paranthropus and Homo. The Pliocene paleoanthropological site of Laetoli in Tanzania provides an opportunity to investigate the relationship between vegetation change and hominin turnover because it encompasses the time period when grass cover was spreading across eastern Africa and because hominin species turnover occurred locally at Laetoli, with Paranthropus aethiopicus in the Upper Ndolanya Beds (UNB) replacing Australopithecus afarensis in the Upper Laetolil Beds (ULB). However, it remains unresolved how the vegetation of the UNB and the ULB differed from each other. To examine differences between the two stratigraphic units, multiple proxies-hypsodonty, mesowear, and stable carbon isotopes of tooth enamel (δ13Cenamel)-are used to infer the diets of large herbivores and compare the dietary guild structure of the large herbivore communities. All three proxies indicate an increase in the abrasiveness and C4-content in the diets of the large herbivores in the UNB relative to those in the ULB. After inferring the diets of species based on all three proxies, the large herbivore community of the UNB had a greater proportion of grazers and a smaller proportion of mixed feeders than in the ULB but maintained a similar proportion of browsers and frugivores. The ULB community has few modern-day analogs, whereas the UNB community is most closely analogous to those in modern African grasslands. Thus, hominin turnover at Laetoli is associated with an increase in grass cover within a woodland-grassland mosaic and is part of a broader transformation of the herbivore community structure.

Edge chipping patterns in posterior teeth of hominins and apes.

Chip scars in fossil teeth are a lasting evidence that bears on human evolution. Chip dimensions in posterior teeth of hominins, apes and white-lipped peccary (Tayassu pecari) are measured from published occlusal images. The results are plotted as D/Dm vs. h/Dm, where h, D and Dm denote indent distance, chip width and mean tooth crown diameter. The hominin species follow a similar pattern where D/Dm monotonically increases up to h/Dm ≈ 0.3. The behavior for the apes is characterized by two phases. In the first, h/Dm monotonically increases up to h/Dm ≈ 0.26 while in the second (h/Dm ≈ 0.26 to 0.42), D/Dm experiences a drastic change in behavior. The interpretation of chip morphology is assisted by results from controlled spherical indentation tests on extracted human molars. This study shows that in addition to the commonly recognized chipping due to cusp loading, a chip may also initiate from the inner wall of the tooth's central fossa. Accordingly, it is suggested that the chipping in hominins generally initiates from a (worn) cusp while that in apes involves cusp loading up to h/Dm ≈ 0.26 and fossa loading thereafter. The behavior for T. pecari is much similar to that of the apes. The fossa chipping is facilitated by a consumption of hard, large-size diet (e.g., plants, roots, barks and nuts) and presence of broad central fossa, conditions that are met in apes. Finally, a simple expression for the critical chipping force Pch due to fossa loading is developed.

Multi-isotope reconstruction of Late Pleistocene large-herbivore biogeography and mobility patterns in Central Europe.

Interpretations of Late Pleistocene hominin adaptative capacities by archaeologists have focused heavily on their exploitation of certain prey and documented contemporary behaviours for these species. However, we cannot assume that animal prey-taxa ecology and ethology were the same in the past as in the present, or were constant over archaeological timescales. Sequential isotope analysis of herbivore teeth has emerged as a particularly powerful method of directly reconstructing diet, ecology and mobility patterns on sub-annual scales. Here, we apply 87Sr/86Sr isotope analysis, in combination with δ18O and δ13C isotope analysis, to sequentially sampled tooth enamel of prevalent herbivore species that populated Europe during the Last Glacial Period, including Rangifer tarandus, Equus sp. and Mammuthus primigenius. Our samples come from two open-air archaeological sites in Central Germany, Königsaue and Breitenbach, associated with Middle Palaeolithic and early Upper Palaeolithic cultures, respectively. We identify potential inter- and intra-species differences in range size and movement through time, contextualised through insights into diet and the wider environment. However, homogeneous bioavailable 87Sr/86Sr across large parts of the study region prevented the identification of specific migration routes. Finally, we discuss the possible influence of large-herbivore behaviour on hominin hunting decisions at the two sites.

Laetoli, Tanzania: Extant terrestrial mollusc faunas shed new light on climate and palaeoecology at a Pliocene hominin site.

Laetoli, Tanzania is one of the most important palaeontological and palaeoanthropological localities in Africa. We report on a survey of the extant terrestrial gastropod faunas of the Laetoli-Endulen area, examine their ecological associations and re-examine the utility of Pliocene fossil molluscs in palaeoenvironmental reconstruction. Standardised collecting at 15 sites yielded 7302 individuals representing 58 mollusc species. Significant dissimilarities were found among the faunas of three broad habitat types: forest, woodland/bushland and open (grassland and scattered, xeric shrubland). Overall, more species were recorded in the woodland/bushland sites than in the forest sites. Open sites were less diverse. Environmental factors contributing most strongly to the separation of habitat types were aridity index and elevation. The results are supplemented with new mollusc data from the Mbulu Plateau south of Lake Eyasi, and compared to the list of species cumulatively recorded from the Ngorongoro area. Some regional variation is apparent and historical factors may explain the absence of some fossil taxa from Laetoli today. Differences in seasonality separated upland forest sites on the Mbulu plateau from those at Lemagurut at Laetoli. Indicator species were identified for each habitat. These included several large-bodied species analogous to the Laetoli Pliocene fossil species that were then used for palaeoenvironmental reconstruction. Based on the estimated aridity index, and adopting the widely used United Nations Environment Programme (UNEP) global climate classification, the four stratigraphic subunits of the Upper Laetolil Beds (3.6-3.85 Ma) would be placed in either the UNEP's Dry Sub-humid or Semi-arid climate classes, whereas the Upper Ndolanya Beds (2.66 Ma) and Lower Laetolil Beds (3.85-<4.36 Ma) would be assigned to the Humid and Semi-arid climate classes respectively. Pliocene precipitation at Laetoli is estimated as 847-965 mm per year, refining previous estimates. This is close or slightly higher than the present mean annual precipitation, and is likely to have corresponded to a mosaic of forest, woodland and bushland within a grassland matrix consistent with other reconstructions.

Janus faced: The co-evolution of war and peace in the human species.

The human species presents a paradox. No other species possesses the propensity to carry out coalitionary lethal attacks on adult conspecifics coupled with the inclination to establish peaceful relations with genetically unrelated groups. What explains this seemingly contradictory feature? Existing perspectives, the "deep roots" and "shallow roots" of war theses, fail to capture the plasticity of human intergroup behaviors, spanning from peaceful cooperation to warfare. By contrast, this article argues that peace and war have both deep roots, and they co-evolved through an incremental process over several million years. On the one hand, humans inherited the propensity for coalitionary lethal violence from their chimpanzee-like ancestor. Specifically, having first inherited the skills to engage in cooperative hunting, they gradually repurposed such capacity to execute coalitionary killings of adult conspecifics and subsequently enhanced it through tech`nological innovations like the use of weapons. On the other hand, they underwent a process of cumulative cultural evolution and, subsequently, of self-domestication which led to heightened cooperative communication and increased prosocial behavior within and between groups. The combination of these two biocultural evolutionary processes-coupled with feedback loop effects between self-domestication and Pleistocene environmental variability-considerably broadened the human intergroup behavioral repertoire, thereby producing the distinctive combination of conflictual and peaceful intergroup relations that characterizes our species. To substantiate this argument, the article synthesizes and integrates the findings from a variety of disciplines, leveraging evidence from evolutionary anthropology, primatology, archeology, paleo-genetics, and paleo-climatology.

Metatarsals and foot phalanges from the Sima de los Huesos Middle Pleistocene site (Atapuerca, Burgos, Spain).

This study provides a complete, updated and illustrated inventory, as well as a comprehensive study, of the metatarsals and foot phalanges (forefoot) recovered from the Middle Pleistocene site of Sima de los Huesos (SH, Atapuerca, Spain) in comparison to other Homo comparative samples, both extant and fossils. This current updated review has established a minimum number of individuals (MNI) of 17, which represent 58.6% of the 29 dental individuals identified within the SH sample. An exclusive or autoapomorphic combination of traits can be recognized within the SH hominin foot sample. A few traits appear primitive or plesiomorphic when compared with earlier Homo individuals and other recent modern humans. There are other metrical and morphological traits that SH hominins and Neandertals have in common that sometimes represent shared derived traits in this evolutionary line, most of which are probably related to robusticity. Furthermore, some exclusive autoapomorphic traits are observed in the SH sample: a very broad first metatarsal, long and broad hallucal proximal foot phalanges and possibly extremely robust lateral distal foot phalanges compared to those of Neandertals and modern humans. In these last traits, the SH metatarsals and pedal phalanges are even more robust than in Neandertals. They are herein named as "hyper-Neandertal" traits, which could suggest a slight gracilization process in this evolutionary line, at least in the hallux toe. Finally, some paleobiological inferences are made in relation to body size (stature and body mass) and some associations are proposed within the SH sample.

Trabecular architecture of the distal femur in extant hominids.

Extant great apes are characterized by a wide range of locomotor, postural and manipulative behaviours that each require the limbs to be used in different ways. In addition to external bone morphology, comparative investigation of trabecular bone, which (re-)models to reflect loads incurred during life, can provide novel insights into bone functional adaptation. Here, we use canonical holistic morphometric analysis (cHMA) to analyse the trabecular morphology in the distal femoral epiphysis of Homo sapiens (n = 26), Gorilla gorilla (n = 14), Pan troglodytes (n = 15) and Pongo sp. (n = 9). We test two predictions: (1) that differing locomotor behaviours will be reflected in differing trabecular architecture of the distal femur across Homo, Pan, Gorilla and Pongo; (2) that trabecular architecture will significantly differ between male and female Gorilla due to their different levels of arboreality but not between male and female Pan or Homo based on previous studies of locomotor behaviours. Results indicate that trabecular architecture differs among extant great apes based on their locomotor repertoires. The relative bone volume and degree of anisotropy patterns found reflect habitual use of extended knee postures during bipedalism in Homo, and habitual use of flexed knee posture during terrestrial and arboreal locomotion in Pan and Gorilla. Trabecular architecture in Pongo is consistent with a highly mobile knee joint that may vary in posture from extension to full flexion. Within Gorilla, trabecular architecture suggests a different loading of knee in extension/flexion between females and males, but no sex differences were found in Pan or Homo, supporting our predictions. Inter- and intra-specific variation in trabecular architecture of distal femur provides a comparative context to interpret knee postures and, in turn, locomotor behaviours in fossil hominins.

Evolution of the throwing shoulder: why apes don't throw well and how that applies to throwing athletes.

BACKGROUND: Humans have unique characteristics making us the only primate that can throw well while most other primates throw predominately underhand with poor speed and accuracy. The purpose of this study is to illuminate the uniquely human characteristics that allow us to throw so well. When treating an injury such as a labral tear or capsule tear, this study hopes the reader can gain a better understanding of the issues that lead to the tear and those that may determine the success of treatment besides the actual repair. METHODS: In addition to a review of scientific and medical literature, information was obtained from interviews and experience with primate veterinarians, anthropologists, archeologists, and professional baseball players. These sources were used to study the connection between evolutionary throwing activities and current sports medicine issues. RESULTS: Arm acceleration requires a functional kinetic chain, rapid motor sequences, and the ability to absorb elastic energy in the shoulder. Successful treatment of the throwing shoulder requires awareness of the shoulder's position in the kinetic chain and correction of defects in the ability to execute the kinetic chain. Some problems in the shoulder could reflect regression to a more primitive anatomy or dyskinesis. Return of performance requires regaining the elasticity in the tissues of the shoulder to temporarily store kinetic energy. For example, tissue remodeling after rotator cuff repair continues for months to years; however, the newly formed tissue lacks the same elasticity of the native tendon. This suggests why throwing performance typically does not return for 7 or more months after repair even though there may be structural integrity at 3-4 months. CONCLUSION: The shoulder has developed uniquely in modern man for the act of throwing. The anatomic deficiencies in primates for throwing provide an illustration of the more subtle changes that a throwing athlete might have that are detrimental to throwing. Nonhuman primates have been unable to demonstrate the kinetic chain sequence for throwing secondary to the lack of neurologic pathways required. Humans are more sophisticated and precise in their movements but lack robusticity in their bone and muscle architecture, seen especially in the human rotator cuff. Successful treatment of a throwing injury requires familiarity with the conditions that cause the injury or affect the rehabilitation process. The return of performance following injury or surgery requires regaining the elasticity in the tissues of the shoulder to temporarily store kinetic energy from the kinetic chain.

On some statistical and cerebral aspects of the limits of working memory capacity in anthropoid primates, with particular reference to Pan and Homo, and their significance for human evolution.

Some comparative ontogenetic data imply that effective working-memory capacity develops in ways that are independent of brain size in humans. These are interpreted better from neuroscientific considerations about the continuing development of neuronal architecture in adolescents and young adults, than from one about gross brain mass which already is reached in childhood. By contrast, working-memory capacity in Pan never develops beyond that of three- or four-year-old children. The phylogenetic divergence begs the question of whether it is any longer plausible to infer from the fossil record, that over the past two million years, an ostensibly gradual increase in endocranial volumes, assigned to the genus Homo, can be correlated in a scientifically-meaningful manner with the gradual evolution of our effective executive working memory. It is argued that whereas Pan's effective working-memory capacity is relatively similar to that of its storage working-memory, our working memory is relatively larger with deeper executive control.

A three-dimensional musculoskeletal model of the pelvis and lower limb of Australopithecus afarensis.

OBJECTIVES: Musculoskeletal modeling is a powerful approach for studying the biomechanics and energetics of locomotion. Australopithecus (A.) afarensis is among the best represented fossil hominins and provides critical information about the evolution of musculoskeletal design and locomotion in the hominin lineage. Here, we develop and evaluate a three-dimensional (3-D) musculoskeletal model of the pelvis and lower limb of A. afarensis for predicting muscle-tendon moment arms and moment-generating capacities across lower limb joint positions encompassing a range of locomotor behaviors. MATERIALS AND METHODS: A 3-D musculoskeletal model of an adult A. afarensis pelvis and lower limb was developed based primarily on the A.L. 288-1 partial skeleton. The model includes geometric representations of bones, joints and 35 muscle-tendon units represented using 43 Hill-type muscle models. Two muscle parameter datasets were created from human and chimpanzee sources. 3-D muscle-tendon moment arms and isometric joint moments were predicted over a wide range of joint positions. RESULTS: Predicted muscle-tendon moment arms generally agreed with skeletal metrics, and corresponded with human and chimpanzee models. Human and chimpanzee-based muscle parameterizations were similar, with some differences in maximum isometric force-producing capabilities. The model is amenable to size scaling from A.L. 288-1 to the larger KSD-VP-1/1, which subsumes a wide range of size variation in A. afarensis. DISCUSSION: This model represents an important tool for studying the integrated function of the neuromusculoskeletal systems in A. afarensis. It is similar to current human and chimpanzee models in musculoskeletal detail, and will permit direct, comparative 3-D simulation studies.

Comparative transcriptomics reveals human-specific cortical features.

The cognitive abilities of humans are distinctive among primates, but their molecular and cellular substrates are poorly understood. We used comparative single-nucleus transcriptomics to analyze samples of the middle temporal gyrus (MTG) from adult humans, chimpanzees, gorillas, rhesus macaques, and common marmosets to understand human-specific features of the neocortex. Human, chimpanzee, and gorilla MTG showed highly similar cell-type composition and laminar organization as well as a large shift in proportions of deep-layer intratelencephalic-projecting neurons compared with macaque and marmoset MTG. Microglia, astrocytes, and oligodendrocytes had more-divergent expression across species compared with neurons or oligodendrocyte precursor cells, and neuronal expression diverged more rapidly on the human lineage. Only a few hundred genes showed human-specific patterning, suggesting that relatively few cellular and molecular changes distinctively define adult human cortical structure.

The promise of great apes as model organisms for understanding the downstream consequences of early life experiences.

Early life experiences have a significant influence on adult health and aging processes in humans. Despite widespread interest in the evolutionary roots of this phenomenon, very little research on this topic has been conducted in humans' closest living relatives, the great apes. The longitudinal data sets that are now available on wild and captive great ape populations hold great promise to clarify the nature, evolutionary function, and mechanisms underlying these connections in species which share key human life history characteristics. Here, we explain features of great ape life history and socioecologies that make them of particular interest for this topic, as well as those that may limit their utility as comparative models; outline the ways in which available data are complementary to and extend the kinds of data that are available for humans; and review what is currently known about the connections among early life experiences, social behavior, and adult physiology and biological fitness in our closest living relatives. We conclude by highlighting key next steps for this emerging area of research.

Exploring the relative influence of raw materials, percussion techniques, and hominin skill levels on the diversity of the early Oldowan assemblages: Insights from the Shungura Formation, Lower Omo Valley, Ethiopia.

The eastern African Oldowan has been documented in multiple raw material contexts and physical environments and displays considerable differences in terms of technological complexity. The relative influence of percussion techniques and raw material quality are central to debates concerning hominin skill levels as a potential driver of change during the period between 2.6 and 2 million-years (Ma). The early Oldowan assemblages from the Shugura Formation play a key role in these debates due to a number of distinctive features, including the small size of the artefacts and poorly controlled flaking. Here we mobilize quantified and replicable experimental data in order to (a) assess the significance of the bipolar technique in the Omo archaeological assemblages and (b) discriminate the respective impact of raw materials, technical choices and knapper skill levels on the unique character of these assemblages. By combining descriptive statistics with regression tree models, our analysis demonstrates knapper skill level to be of minimal importance in this context for the production of sharp-edged flakes. The absence of a link between skill and knapping success reflects the combined effect of raw material constraints, the frequent use of the bipolar technique, and relatively simple technical objectives. Our analysis confirms the key role played by local environmental conditions in the unique appearance of the Shungura assemblages, a relationship which has been frequently suggested but never demonstrated. Beyond the operational and sensorimotor skills considered in most studies, we suggest that the diversity of early Oldowan assemblages should be better sought in the cognitive abilities developed by early toolmakers as a response to landscape learning and use, two elements of early human evolution that remain largely unexplored.

The aorta in humans and African great apes, and cardiac output and metabolic levels in human evolution.

Humans have a larger energy budget than great apes, allowing the combination of the metabolically expensive traits that define our life history. This budget is ultimately related to the cardiac output, the product of the blood pumped from the ventricle and the number of heart beats per minute, a measure of the blood available for the whole organism physiological activity. To show the relationship between cardiac output and energy expenditure in hominid evolution, we study a surrogate measure of cardiac output, the aortic root diameter, in humans and great apes. When compared to gorillas and chimpanzees, humans present an increased body mass adjusted aortic root diameter. We also use data from the literature to show that over the human lifespan, cardiac output and total energy expenditure follow almost identical trajectories, with a marked increase during the period of brain growth, and a plateau during most of the adult life. The limited variation of adjusted cardiac output with sex, age and physical activity supports the compensation model of energy expenditure in humans. Finally, we present a first study of cardiac output in the skeleton through the study of the aortic impression in the vertebral bodies of the spine. It is absent in great apes, and present in humans and Neanderthals, large-brained hominins with an extended life cycle. An increased adjusted cardiac output, underlying higher total energy expenditure, would have been a key process in human evolution.

The evolution of hominoid locomotor versatility: Evidence from Moroto, a 21 Ma site in Uganda.

Living hominoids are distinguished by upright torsos and versatile locomotion. It is hypothesized that these features evolved for feeding on fruit from terminal branches in forests. To investigate the evolutionary context of hominoid adaptive origins, we analyzed multiple paleoenvironmental proxies in conjunction with hominoid fossils from the Moroto II site in Uganda. The data indicate seasonally dry woodlands with the earliest evidence of abundant C4 grasses in Africa based on a confirmed age of 21 million years ago (Ma). We demonstrate that the leaf-eating hominoid Morotopithecus consumed water-stressed vegetation, and postcrania from the site indicate ape-like locomotor adaptations. These findings suggest that the origin of hominoid locomotor versatility is associated with foraging on leaves in heterogeneous, open woodlands rather than forests.

To copy or not to copy? That is the question! From chimpanzees to the foundation of human technological culture.

A prerequisite for copying innovative behaviour faithfully is the capacity of observers' brains, regarded as 'hierarchically mechanistic minds', to overcome cognitive 'surprisal' (see 2.), by maximising the evidence for their internal models, through active inference. Unlike modern humans, chimpanzees and other great apes show considerable limitations in their ability, or 'Zone of Bounded Surprisal', to overcome cognitive surprisal induced by innovative or unorthodox behaviour that rarely, therefore, is copied precisely or accurately. Most can copy adequately what is within their phenotypically habitual behavioural repertoire, in which technology plays scant part. Widespread intra- and intergenerational social transmission of complex technological innovations is not a hall-mark of great-ape taxa. 3 Ma, precursors of the genus Homo made stone artefacts, and stone-flaking likely was habitual before 2 Ma. After that time, early Homo erectus has left traces of technological innovations, though faithful copying of these and their intra- and intergenerational social transmission were rare before 1 Ma. This likely owed to a cerebral infrastructure of interconnected neuronal systems more limited than ours. Brains were smaller in size than ours, and cerebral neuronal systems ceased to develop when early Homo erectus attained full adult maturity by the mid-teen years, whereas its development continues until our mid-twenties nowadays. Pleistocene Homo underwent remarkable evolutionary adaptation of neurobiological propensities, and cerebral aspects are discussed that, it is proposed here, plausibly, were fundamental for faithful copying, which underpinned social transmission of technologies, cumulative learning, and culture. Here, observers' responses to an innovation are more important for ensuring its transmission than is an innovator's production of it, because, by themselves, the minimal cognitive prerequisites that are needed for encoding and assimilating innovations are insufficient for practical outcomes to accumulate and spread intra- and intergenerationally.

Evolutionary ecology of language origins through confrontational scavenging.

A dynamic model and an agent-based simulation model implementing the assumptions of the confrontational scavenging hypothesis on early protolanguage as an adaptive response of Homo erectus to gradual change in their habitat has been developed and studied. The core assumptions of the hypothesis and the model scenario are the pre-adaptation of our ancestors to occupy the ecological niche that they constructed for themselves by having evolved displaced communication and a rudimentary tool manufacture, two features allowing them to use a new, concentrated and abundant resource-megafauna carrion-on the savannahs replacing arboreal habitats owing to the drying climate of East Africa at about 2 Ma. The shift in diet required coordinated cooperation by the hominin scavengers confronted with concurrent predators. Power scavenging compelled displaced symbolic communication featuring a limited semantic range; syntax was not yet required. We show that phenotypic evolution on the accuracy of information transfer between cooperating hominins is a necessary and sufficient condition for the population of agents to survive the diet shift. Both the individual and the group fitness of the hominin horde increased with the accuracy of their protolanguage, with decreasing time allocated to foraging and thus more time left for culture. This article is part of the theme issue 'Human socio-cultural evolution in light of evolutionary transitions'.

Language as a tool for social bonding: evidence from wild chimpanzee gestural, vocal and bimodal signals.

The evolution of language has fascinated anthropologists, psychologists and biologists for centuries, seeking to infer language origins from the communication of primates, our closest living relatives. Capacity for intentional signalling is a key feature of transition to language in our hominin ancestors, facilitating complex social dynamics in complex social groups. However whether vocal, gestural and bimodal signals are differentiated according to intentional use and hence complex sociality has not been studied, making unclear the modality of language evolution. We addressed this question in wild chimpanzees. We found that larger social network size was associated with a larger network of gestural but not vocal or bimodal signals. Response waiting was more common in association with gestures than vocalizations, but elaborations were more common in vocal than gestural or bimodal signals. Overall, chimpanzees were more likely to manage weak social bonds through vocalizations, whereas strong social bonds were managed through gestures and bimodal signals. However, when social bonds were weak, gestures accompanied by response waiting were more likely to elicit approaches than vocalizations accompanied by elaboration, which elicited avoidance. This suggests that gestures were the primary modality of language evolution and that the use of more sophisticated gestural signalling led to evolution of complex social groups of hominin ancestors. This article is part of the theme issue 'Cognition, communication and social bonds in primates'.