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Syntax is widely considered the feature that most decisively sets human language apart from other natural communication systems. Animal vocalisations are generally considered to be holistic with few examples of utterances meaning something other than the sum of their parts. Previously, we have shown that male putty-nosed monkeys produce call series consisting of two call types in response to different events. They can also be combined into short sequences that convey a different message from those conveyed by either call type alone. Here, we investigate whether 'pyow-hack' sequences are compositional in that the individual calls contribute to their overall meaning. However, the monkeys behaved as if they perceived the sequence as an idiomatic expression rather than decoding the sequence. Nonetheless, while this communication system lacks the generative power of syntax it enables callers to increase the number of messages that can be conveyed by a small and innate call repertoire. [ABSTRACT FROM AUTHOR]

Data are missing on the diversity of Plasmodium spp. infecting apes that live in their natural habitat, with limited possibility of human-mosquito-ape exchange. We surveyed Plasmodium spp. diversity in wild chimpanzees living in an undisturbed tropical rainforest habitat and found 5 species: P. malariae, P. vivax, P. ovale, P. reichenowi, and P. gaboni. [ABSTRACT FROM AUTHOR]

The article presents an abstract of the paper "Predation and Primate Cognitive Evolution," by K. Zuberbühler, at the Twenty-First Congress of the International Primatological Society to be held in Entebbe, Uganda, on June 25-30, 2006.

The article presents the abstract of the paper "Spatial Memory of Fruiting States of Rain Forest Trees in Wild Ranging Mangabeys," by K.R. Janmaat and colleagues, to be presented at the 21st Congress of the International Primatological Society in Entebbe, Uganda from June 25-30, 2006.

The article presents an abstract of the paper "Do mangabeys take past weather into account when they revisit ficus trees with fruit," by K.R. Janmaat, R.W. Byrne and K. Zuberbühler, to be presented at the 21st Congress of the International Society of Primatological Society in Entebbe, Uganda on June 25-30, 2006.

Abstract Background Non-human primate communication is thought to be fundamentally different from human speech, mainly due to vast differences in vocal control. The lack of these abilities in non-human primates is especially striking if compared to some marine mammals and bird species, which has generated somewhat of an evolutionary conundrum. What are the biological roots and underlying evolutionary pressures of the human ability to voluntarily control sound production and learn the vocal utterances of others? One hypothesis is that this capacity has evolved gradually in humans from an ancestral stage that resembled the vocal behavior of modern primates. Support for this has come from studies that have documented limited vocal flexibility and convergence in different primate species, typically in calls used during social interactions. The mechanisms underlying these patterns, however, are currently unknown. Specifically, it has been difficult to rule out explanations based on genetic relatedness, suggesting that such vocal flexibility may not be the result of social learning. Results To address this point, we compared the degree of acoustic similarity of contact calls in free-ranging Campbell's monkeys as a function of their social bonds and genetic relatedness. We calculated three different indices to compare the similarities between the calls' frequency contours, the duration of grooming interactions and the microsatellite-based genetic relatedness between partners. We found a significantly positive relation between bond strength and acoustic similarity that was independent of genetic relatedness. Conclusion Genetic factors determine the general species-specific call repertoire of a primate species, while social factors can influence the fine structure of some the call types. The finding is in line with the more general hypothesis that human speech has evolved gradually from earlier primate-like vocal communication.

The article presents an abstract of the paper "Functionally Referential Calls in Chimpanzees," by K. Zuberbühler and K. Slocombe, to be presented at the 21st Congress of the International Society of Primatological Society in Entebbe, Uganda on June 25-30, 2006.

The ability to coordinate actions is of vital importance for group-living animals, particularly in relation to travel. Groups can only remain cohesive if members possess a cooperative mechanism to overcome differences in individual priorities and social power when coordinating departures. To better understand how hominids achieve spatio-temporally coordinated group movements, we investigated vocally initiated group departures in three habituated groups of western gorillas ( Gorilla gorilla ) in the Central African Republic. The large sexual dimorphism of gorillas has led to the untested assumption that the silverback males are the sole decision-makers in gorilla groups, although there are also observations that suggest otherwise. To address this, we analysed the direction and timing of group departures and found that high-ranking individuals (silverbacks and high-ranking females) were more successful in indicating the direction of future travel than others, but that the timing of departure was the apparent result of a cumulative vocal voting process among all adult group members. Our findings illustrate that even in species with a large sexual size dimorphism, travel decisions can be taken collectively via a consensus-building process.

Parsing signals from noise is a general problem for signallers and recipients, and for researchers studying communicative systems. Substantial efforts have been invested in comparing how other species encode information and meaning, and how signalling is structured. However, research depends on identifying and discriminating signals that represent meaningful units of analysis. Early approaches to defining signal repertoires applied top-down approaches, classifying cases into predefined signal types. Recently, more labour-intensive methods have taken a bottom-up approach describing detailed features of each signal and clustering cases based on patterns of similarity in multi-dimensional feature-space that were previously undetectable. Nevertheless, it remains essential to assess whether the resulting repertoires are composed of relevant units from the perspective of the species using them, and redefining repertoires when additional data become available. In this paper we provide a framework that takes data from the largest set of wild chimpanzee (Pan troglodytes) gestures currently available, splitting gesture types at a fine scale based on modifying features of gesture expression using latent class analysis (a model-based cluster detection algorithm for categorical variables), and then determining whether this splitting process reduces uncertainty about the goal or community of the gesture. Our method allows different features of interest to be incorporated into the splitting process, providing substantial future flexibility across, for example, species, populations, and levels of signal granularity. Doing so, we provide a powerful tool allowing researchers interested in gestural communication to establish repertoires of relevant units for subsequent analyses within and between systems of communication., (© 2024. The Author(s).)

Non-human primates generally lack the ability to learn new call structures or to substantially modify existing ones, suggesting that callers need alternative mechanisms to convey information. One way to escape the constraints of limited vocal control is by assembling calls into variable sequences, as has been documented in various animal species. Here, we were interested in the flexibility with which different calls might be assembled in a species known for its meaningful call order, putty-nosed monkeys (Cercopithecus nictitans). Since most information comes from studies conducted at Gashaka Gumti National Park (Nigeria), we tested two further populations in the Nouabalé-Ndoki National Park (Republic of the Congo) and Taï National Park (Côte d'Ivoire) in how males responded to common threats, leopards, and crowned eagles. As predicted, callers produced the same basic call types as seen elsewhere-long 'pyow', short 'pyow' ('kek'), 'hack'-but populations differed in how males assembled calls. To leopards, males from both populations started with 'pyows' and 'keks', with occasional hacks later, as already reported from Gashaka. To crowned eagle, however, Nouabalé-Ndoki males consistently initiated their responses with 'pyows', whereas neither Taï nor Gashaka males ever did, demonstrating that nonhuman primates have some control over sequence production. We discuss possible mechanisms to account for the population differences, predation pressure, and male-male competition, and address implications for linguistic theories of animal call order, notably the Urgency and Informativity Principles., (© 2024. The Author(s), under exclusive licence to Japan Monkey Centre.)

As one of the most specific, yet most diverse of human behaviors, language is shaped by both genomic and extra-genomic evolution. Sharing methods and models between these modes of evolution has significantly advanced our understanding of language and inspired generalized theories of its evolution. Progress is hampered, however, by the fact that the extra-genomic evolution of languages, i.e. linguistic evolution, maps only partially to other forms of evolution. Contrasting it with the biological evolution of eukaryotes and the cultural evolution of technology as the best understood models, we show that linguistic evolution is special by yielding a stationary dynamic rather than stable solutions, and that this dynamic allows the use of language change for social differentiation while maintaining its global adaptiveness. Linguistic evolution furthermore differs from technological evolution by requiring vertical transmission, allowing the reconstruction of phylogenies; and it differs from eukaryotic biological evolution by foregoing a genotype vs phenotype distinction, allowing deliberate and biased change. Recognising these differences will improve our empirical tools and open new avenues for analyzing how linguistic, cultural, and biological evolution interacted with each other when language emerged in the hominin lineage. Importantly, our framework will help to cope with unprecedented scientific and ethical challenges that presently arise from how rapid cultural evolution impacts language, most urgently from interventional clinical tools for language disorders, potential epigenetic effects of technology on language, artificial intelligence and linguistic communicators, and global losses of linguistic diversity and identity. Beyond language, the distinctions made here allow identifying variation in other forms of biological and cultural evolution, developing new perspectives for empirical research., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024. Published by Elsevier B.V.)

Humans regularly engage in efficient communicative conversations, which serve to socially align individuals 1 . In conversations, we take fast-paced turns using a human-universal structure of deploying and receiving signals which shows consistent timing across cultures 2 . We report here that chimpanzees also engage in rapid signal-to-signal turn-taking during face-to-face gestural exchanges with a similar average latency between turns to that of human conversation. This correspondence between human and chimpanzee face-to-face communication points to shared underlying rules in communication. These structures could be derived from shared ancestral mechanisms or convergent strategies that enhance coordinated interactions or manage competition for communicative 'space'., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2024 The Authors. Published by Elsevier Inc. All rights reserved.)

Wild chimpanzees consume a variety of plants to meet their dietary needs and maintain wellbeing. While some plants have obvious value, others are nutritionally poor and/or contain bioactive toxins which make ingestion costly. In some cases, these nutrient-poor resources are speculated to be medicinal, thought to help individuals combat illness. In this study, we observed two habituated chimpanzee communities living in the Budongo Forest, Uganda, and collected 17 botanical samples associated with putative self-medication behaviors (e.g., bark feeding, dead wood eating, and pith-stripping) or events (e.g., when consumer had elevated parasite load, abnormal urinalysis, or injury). In total, we selected plant parts from 13 species (nine trees and four herbaceous plants). Three extracts of different polarities were produced from each sample using n-hexane, ethyl acetate, and methanol/water (9/1, v/v) and introduced to antibacterial and anti-inflammatory in vitro models. Extracts were evaluated for growth inhibition against a panel of multidrug-resistant clinical isolates of bacteria, including ESKAPE strains and cyclooxygenase-2 (COX-2) inhibition activity. Pharmacological results suggest that Budongo chimpanzees consume several species with potent medicinal properties. In the antibacterial library screen, 45 out of 53 extracts (88%) exhibited ≥40% inhibition at a concentration of 256 μg/mL. Of these active extracts, 41 (91%) showed activity at ≤256μg/mL in subsequent dose-response antibacterial experiments. The strongest antibacterial activity was achieved by the n-hexane extract of Alstonia boonei dead wood against Staphylococcus aureus (IC50: 16 μg/mL; MIC: 32 μg/mL) and Enterococcus faecium (IC50: 16 μg/mL; MIC: >256 μg/mL) and by the methanol-water extract of Khaya anthotheca bark and resin against E. faecium (IC50: 16 μg/mL; MIC: 32 μg/mL) and pathogenic Escherichia coli (IC50: 16 μg/mL; MIC: 256 μg/mL). We observed ingestion of both these species by highly parasitized individuals. K. anthotheca bark and resin were also targeted by individuals with indicators of infection and injuries. All plant species negatively affected growth of E. coli. In the anti-inflammatory COX-2 inhibition library screen, 17 out of 51 tested extracts (33%) showed ≥50% COX-2 inhibition at a concentration of 5 μg/mL. Several extracts also exhibited anti-inflammatory effects in COX-2 dose-response experiments. The K. anthotheca bark and resin methanol-water extract showed the most potent effects (IC50: 0.55 μg/mL), followed by the fern Christella parasitica methanol-water extract (IC50: 0.81 μg/mL). This fern species was consumed by an injured individual, a feeding behavior documented only once before in this population. These results, integrated with associated observations from eight months of behavioral data, provide further evidence for the presence of self-medicative resources in wild chimpanzee diets. This study addresses the challenge of distinguishing preventative medicinal food consumption from therapeutic self-medication by integrating pharmacological, observational, and health monitoring data-an essential interdisciplinary approach for advancing the field of zoopharmacognosy., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Freymann et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

When conversing, humans instantaneously predict meaning from fragmentary and ambiguous mspeech, long before utterance completion. They do this by integrating priors (initial assumptions about the world) with contextual evidence to rapidly decide on the most likely meaning. One powerful prior is attentional preference for agents, which biases sentence processing but universally so only if agents are animate. Here, we investigate the evolutionary origins of this preference, by allowing chimpanzees, gorillas, orangutans, human children, and adults to freely choose between agents and patients in still images, following video clips depicting their dyadic interaction. All participants preferred animate (and occasionally inanimate) agents, although the effect was attenuated if patients were also animate. The findings suggest that a preference for animate agents evolved before language and is not reducible to simple perceptual biases. To conclude, both humans and great apes prefer animate agents in decision tasks, echoing a universal prior in human language processing., Competing Interests: The authors declare no competing interests., (© 2024 The Author(s).)

Identifying novel medicinal resources in chimpanzee diets has historically presented challenges, requiring extensive behavioral data collection and health monitoring, accompanied by expensive pharmacological analyses. When putative therapeutic self-medicative behaviors are observed, these events are often considered isolated occurrences, with little attention paid to other resources ingested in combination. For chimpanzees, medicinal resource combinations could play an important role in maintaining well-being by tackling different symptoms of an illness, chemically strengthening efficacy of a treatment, or providing prophylactic compounds that prevent future ailments. We call this concept the self-medicative resource combination hypothesis. However, a dearth of methodological approaches for holistically investigating primate feeding ecology has limited our ability to identify nonrandom resource combinations and explore potential synergistic relationships between medicinal resource candidates. Here we present two analytical tools that test such a hypothesis and demonstrate these approaches on feeding data from the Sonso chimpanzee community in Budongo Forest, Uganda. Using 4 months of data, we establish that both collocation and APRIORI analyses are effective exploratory tools for identifying binary combinations, and that APRIORI is effective for multi-item rule associations. We then compare outputs from both methods, finding up to 60% agreement, and propose APRIORI as more effective for studies requiring control over confidence intervals and those investigating nonrandom associations between more than two resources. These analytical tools, which can be extrapolated across the animal kingdom, can provide a cost-effective and efficient method for targeting resources for further pharmacological investigation, potentially aiding in the discovery of novel medicines., (© 2024 The Authors. American Journal of Primatology published by Wiley Periodicals LLC.)

Proposed mechanisms of zoonotic virus spillover often posit that wildlife transmission and amplification precede human outbreaks. Between 2006 and 2012, the palm Raphia farinifera, a rich source of dietary minerals for wildlife, was nearly extirpated from Budongo Forest, Uganda. Since then, chimpanzees, black-and-white colobus, and red duiker were observed feeding on bat guano, a behavior not previously observed. Here we show that guano consumption may be a response to dietary mineral scarcity and may expose wildlife to bat-borne viruses. Videos from 2017-2019 recorded 839 instances of guano consumption by the aforementioned species. Nutritional analysis of the guano revealed high concentrations of sodium, potassium, magnesium and phosphorus. Metagenomic analyses of the guano identified 27 eukaryotic viruses, including a novel betacoronavirus. Our findings illustrate how "upstream" drivers such as socioeconomics and resource extraction can initiate elaborate chains of causation, ultimately increasing virus spillover risk., (© 2024. The Author(s).)

Primate social organizations, or grouping patterns, vary significantly across species. Behavioral strategies that allow for flexibility in grouping patterns offer a means to reduce the costs of group living. Chimpanzees (Pan troglodytes) have a fission-fusion social system in which temporary subgroups ("parties") change in composition because of local socio-ecological conditions. Notably, western chimpanzees (P. t. verus) are described as showing a higher degree of bisexual bonding and association than eastern chimpanzees, and eastern female chimpanzees (P. t. schweinfurthii) are thought to be more solitary than western female chimpanzees. However, reported comparisons in sociality currently depend on a small number of study groups, particularly in western chimpanzees, and variation in methods. The inclusion of additional communities and direct comparison using the same methods are essential to assess whether reported subspecies differences in sociality hold in this behaviorally heterogeneous species. We explored whether sociality differs between two communities of chimpanzees using the same motion-triggered camera technology and definitions of social measures. We compare party size and composition (party type, sex ratio) between the western Gahtoy community in the Nimba Mountains (Guinea) and the eastern Waibira community in the Budongo Forest (Uganda). Once potential competition for resources such as food and mating opportunities were controlled for, subspecies did not substantially influence the number of individuals in a party. We found a higher sex-ratio, indicating more males in a party, in Waibira; this pattern was driven by a greater likelihood in Gahtoy to be in all-female parties. This finding is the opposite of what was expected for eastern chimpanzees, where female-only parties are predicted to be more common. Our results highlight the flexibility in chimpanzee sociality, and caution against subspecies level generalizations., (© 2024 The Authors. American Journal of Primatology published by Wiley Periodicals LLC.)

The social complexity hypothesis for the evolution of communication posits that complex social environments require greater communication complexity for individuals to effectively manage their relationships. We examined how different socially uncertain contexts, reflecting an increased level of social complexity, relate to variation in signalling within and between two species, which display varying levels of fission-fusion dynamics (sympatric-living chimpanzees and sooty mangabeys, Taï National Park, Ivory Coast). Combined signalling may improve message efficacy, notably when involving different perception channels, thus may increase in moments of high social uncertainty. We examined the probability of individuals to emit no signal, single or multisensory or combined (complex) signals, during social approaches which resulted in non-agonistic outcomes. In both species, individuals were more likely to use more combined and multisensory signals in post-conflict approaches with an opponent than in other contexts. The clearest impact of social uncertainty on signalling complexity was observed during chimpanzee fusions, where the likelihood of using complex signals tripled relative to other contexts. Overall, chimpanzees used more multisensory signals than mangabeys. Social uncertainty may shape detected species differences in variation in signalling complexity, thereby supporting the hypothesis that social complexity, particularly associated with high fission-fusion dynamics, promotes signalling complexity., Competing Interests: We declare we have no competing interests., (© 2023 The Authors.)

Primate alarm calls are mainly hardwired but individuals need to adapt their calling behaviours according to the situation. Such learning necessitates recognising locally relevant dangers and may take place via their own experience or by observing others. To investigate monkeys alarm calling behaviour, we carried out a field experiment in which we exposed juvenile vervet monkeys to unfamiliar raptor models in the presence of audiences that differed in experience and reliability. We used audience age as a proxy for experience and relatedness as a proxy for reliability, while quantifying audience reactions to the models. We found a negative correlation between alarm call production and callers' age. Adults never alarm called, compared to juveniles. We found no overall effect of audience composition and size, with juveniles calling more when with siblings than mothers or unrelated individuals. Finally, concerning audience reactions to the models, we observed juveniles remained silent with vigilant mothers and only alarm called with ignoring mothers, whereas we observed the opposite for siblings: juveniles remained silent with ignoring siblings and called with vigilant siblings. Despite the small sample size, juvenile vervet monkeys, confronted with unfamiliar and potentially dangerous raptors, seem to rely on others to decide whether to alarm call, demonstrating that the choice of the model may play an important key role in the ontogeny of primate alarm call behaviour., (© 2023. The Author(s).)

Social learning is beneficial in almost every domain of a social animal's life, but it is particularly important in the context of predation and foraging. In both contexts, social animals tend to produce acoustically distinct vocalizations, alarms, and food calls, which have remained somewhat of an evolutionary conundrum as they appear to be costly for the signaller. Here, we investigated the hypothesis that food calls function to direct others toward novel food items, using a playback experiment on a group of chimpanzees. We showed chimpanzees novel (plausibly edible) items while simultaneously playing either conspecific food calls or acoustically similar greeting calls as a control. We found that individuals responded by staying longer near items previously associated with food calls even in the absence of these vocalizations, and peered more at these items compared with the control items, provided no conspecifics were nearby. We also found that once chimpanzees had access to both item types, they interacted more with the one previously associated with food calls than the control items. However, we found no evidence of social learning per se. Given these effects, we propose that food calls may gate and thus facilitate social learning by directing listeners' attention to new feeding opportunities, which if integrated with additional cues could ultimately lead to new food preferences within social groups., (© 2023 The Authors. American Journal of Primatology published by Wiley Periodicals LLC.)

The alarm calls of nonhuman primates are occasionally cited as functionally equivalent to lexical word meaning in human language. Recently, however, it has become increasingly unlikely that one-to-one relations between alarm call structures and predator categories are the default, mainly because many call types are produced in multiple contexts, requiring more complex notions of meaning. For example, male vervet monkeys produce the same alarm calls during encounters with terrestrial predators and neighbouring groups, suggesting that recipients require additional information to attribute meaning to the calls. We empirically tested the hypothesis that vervet monkeys take contextual information into account when responding to each other's alarm calls. In playback experiments, we exposed subjects to recordings of male alarm barks during actual intergroup encounters (predator unlikely) or when there was no intergroup encounter (predator likely). Subjects responded more strongly in the no intergroup encounter situations, typically associated with discovering a hiding predator, measured in terms of startle responses, vigilance behaviour and gazing towards the presumed caller. We discuss the significance of using contextual information for meaning attribution in nonhuman primate communication., (© 2023. The Author(s).)

Through syntax, i.e., the combination of words into larger phrases, language can express a limitless number of messages. Data in great apes, our closest-living relatives, are central to the reconstruction of syntax's phylogenetic origins, yet are currently lacking. Here, we provide evidence for syntactic-like structuring in chimpanzee communication. Chimpanzees produce "alarm-huus" when surprised and "waa-barks" when potentially recruiting conspecifics during aggression or hunting. Anecdotal data suggested chimpanzees combine these calls specifically when encountering snakes. Using snake presentations, we confirm call combinations are produced when individuals encounter snakes and find that more individuals join the caller after hearing the combination. To test the meaning-bearing nature of the call combination, we use playbacks of artificially-constructed call combinations and both independent calls. Chimpanzees react most strongly to call combinations, showing longer looking responses, compared with both independent calls. We propose the "alarm-huu + waa-bark" represents a compositional syntactic-like structure, where the meaning of the call combination is derived from the meaning of its parts. Our work suggests that compositional structures may not have evolved de novo in the human lineage, but that the cognitive building-blocks facilitating syntax may have been present in our last common ancestor with chimpanzees., (© 2023. The Author(s).)

While cases of interspecies grooming have been reported in primates, no comprehensive cross-site review has been published about this behavior in great apes. Only a few recorded observations of interspecies grooming events between chimpanzees and other primate species have been reported in the wild, all of which have thus far been in Uganda. Here, we review all interspecies grooming events recorded for the Sonso community chimpanzees in Budongo Forest Reserve, Uganda, adding five new observations to the single, previously reported event from this community. A new case of interspecies play involving three juvenile male chimpanzees and a red-tailed monkey is also detailed. All events took place between 1993 and 2021. In all of the six interspecific grooming events from Budongo, the 'groomer' was a female chimpanzee between the ages of 4-6 years, and the 'recipient' was a member of the genus Cercopithecus. In five of these events, chimpanzee groomers played with the tail of their interspecific grooming partners, and except for one case, initiated the interaction. In three cases, chimpanzee groomers smelled their fingers after touching distinct parts of the receiver's body. While a single function of chimpanzee interspecies grooming remains difficult to determine from these results, our review outlines and assesses some hypotheses for the general function of this behavior, as well as some of the costs and benefits for both the chimpanzee groomers and their sympatric interspecific receivers. As allogrooming is a universal behavior in chimpanzees, investigating the ultimate and proximate drivers of chimpanzee interspecies grooming may reveal further functions of allogrooming in our closest living relatives, and help us to better understand how chimpanzees distinguish between affiliative and agonistic species and contexts., (© 2023. The Author(s).)

Dialects are a cultural property of animal communication previously described in the signals of several animal species. While dialects have predominantly been described in vocal signals, chimpanzee leaf-clipping and other 'leaf-modifying' gestures, used across chimpanzee and bonobo communities, have been suggested as a candidate for cultural variation in gestural communication. Here we combine direct observation with archaeological techniques to compare the form and use of leaf-modifying gestures in two neighbouring communities of East African chimpanzees. We found that while both communities used multiple forms, primarily within sexual solicitation, they showed a strong preference for a single, different gesture form. The observed variation in form preference between these neighbouring communities within the same context suggests that these differences are, at least in part, socially derived. Our results highlight an unexplored source of variation and flexibility in gestural communication, opening the door for future research to explore socially derived dialects in non-vocal communication., (© 2023. The Author(s).)

Abstract: Primates understand the meaning of their own and other species' alarm calls, but little is known about how they acquire such knowledge. Here, we combined direct behavioural observations with playback experiments to investigate two key processes underlying vocal development: comprehension and usage. Especifically, we studied the development of con- and heterospecific alarm call recognition in free-ranging sooty mangabeys, Cercocebus atys , across three age groups: young juveniles (1-2y), old juveniles (3-4y) and adults (> 5y). We observed that, during natural predator encounters, juveniles alarm called to a significantly wider range of species than adults, with evidence of refinement during the first four years of life. In the experiments, we exposed subjects to leopard, eagle and snake alarm calls given by other group members or sympatric Diana monkeys. We found that young juveniles' locomotor and vocal responses were least appropriate and that they engaged in more social referencing (look at adults when hearing an alarm call) than older individuals, suggesting that vocal competence is obtained via social learning. In conclusion, our results suggest that alarm call comprehension is socially learned during the juvenile stage, with comprehension preceding appropriate usage but no difference between learning their own or other species' alarm calls., Significance Statement: Under natural conditions, animals do not just interact with members of their own species, but usually operate in a network of associated species. However, ontogenetic research on primate communication frequently ignores this significant element. We studied the development of con- and heterospecific alarm call recognition in wild sooty mangabeys. We found that communicative competence was acquired during the juvenile stages, with alarm call comprehension learning preceding appropriate vocal usage and with no clear difference in learning of con- and heterospecific signals. We also found that, during early stages of life, social referencing, a proactive form of social learning, was key in the acquisition of competent alarm call behaviour. Our results show that primates equally learn to interpret alarm calls from their own and other species during their early stages of life and that this learning process is refined as the animals mature., Supplementary Information: The online version contains supplementary material available at 10.1007/s00265-023-03318-6., Competing Interests: Competing interestsThe authors declare no competing interests., (© The Author(s) 2023.)

Human economic decision-making sometimes appears to be irrational. Partly, this is due to cognitive biases that can lead to suboptimal economic choices and context-dependent risk-preferences. A pertinent question is whether such biases are part of our evolutionary heritage or whether they are culturally acquired. To address this, we tested gorillas (Gorilla gorilla gorilla) and orang-utans (Pongo abelii) with two risk-assessment experiments that differed in how risk was presented. For both experiments, we found that subjects increased their preferences for the risky options as their expected gains increased, showing basic understanding of reward contingencies and rational decision-making. However, we also found consistent differences in risk proneness between the two experiments, as subjects were risk-neutral in one experiment and risk-prone in the other. We concluded that gorillas and orang-utans are economically rational but that their decisions can interact with pre-existing cognitive biases which modulates their risk-preference in context-dependent ways, explaining the variability of their risk-preference in previous literature., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2022 Lacombe et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Associating with kin provides individual benefits but requires that these relationships be detectable. In humans, facial phenotype matching might help assess paternity; however, evidence for it is mixed. In chimpanzees, concealing visual cues of paternity may be beneficial due to their promiscuous mating system and the considerable risk of infanticide by males. On the other hand, detecting kin can also aid chimpanzees in avoiding inbreeding and in forming alliances that improve kin-mediated fitness. Although previous studies assessing relatedness based on facial resemblance in chimpanzees exist, they used images of captive populations in whom selection pressures and reproductive opportunities are controlled and only assessed maternity or paternity of adult offspring. In natural populations, the chances of infanticide are highest during early infancy, suggesting that young infants would benefit most from paternity concealment, whereas adults and subadults would benefit from the detection of all types of kin, including half-siblings. In our experiment, we conducted an online study with human participants, in which they had to assess the relatedness of chimpanzees based on facial similarity. To address previous methodological constraints, we used chimpanzee images across all ages, as well as maternal and paternal half-siblings. We found that kin status was detected above chance across all relatedness categories, with easier kin detection of father-offspring pairs, females, and older chimpanzees. Together, these findings support the existence of paternity confusion in infant chimpanzees and provide a possible mechanism for incest avoidance and kin-based social alliances in older individuals. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

Spoken language, as we have it, requires specific capacities-at its most basic advanced vocal control and complex social cognition. In humans, vocal control is the basis for speech, achieved through coordinated interactions of larynx activity and rapid changes in vocal tract configurations. Most likely, speech evolved in response to early humans perceiving reality in increasingly complex ways, to the effect that primate-like signaling became unsustainable as a sole communication device. However, in what ways did and do humans see the world in more complex ways compared to other species? Although animal signals can refer to external events, in contrast to humans, they usually refer to the agents only, sometimes in compositional ways, but never together with patients. It may be difficult for animals to comprehend events as part of larger social scripts, with antecedent causes and future consequences, which are more typically tie the patient into the event. Human brain enlargement over the last million years probably has provided the cognitive resources to represent social interactions as part of bigger social scripts, which enabled humans to go beyond an agent-focus to refer to agent-patient relations, the likely foundation for the evolution of grammar. This article is categorized under: Cognitive Biology > Evolutionary Roots of Cognition Linguistics > Evolution of Language Psychology > Comparative., (© 2022 The Authors. WIREs Cognitive Science published by Wiley Periodicals LLC.)

Individuals of social species experience competitive costs and social benefits of group living. Substantial flexibility in humans' social structure and the combination of different types of social structure with fission-fusion dynamics allow us to live in extremely large groups-overcoming some of the costs of group living while capitalizing on the benefits. Non-human species also show a range of social strategies to deal with this trade-off. Chimpanzees are an archetypical fission-fusion species, using dynamic changes in day-to-day association to moderate the costs of within-group competition. Using 4 years of association data from two neighbouring communities of East African chimpanzees ( Pan troglodytes schweinfurthii ), we describe an unexplored level of flexibility in chimpanzee social structure. We show that males from the larger Waibira community ( N = 24-31) exhibited additional structural levels of semi-stable core-periphery society, while males from the smaller Sonso community ( N = 10-13) did not. This novel core-periphery pattern adds to previous results describing alternative modular social structure in other large communities of chimpanzees. Our data support the hypothesis that chimpanzees can incorporate a range of strategies in addition to fission-fusion to overcome costs of social living, and that their social structures may be closer to that of modern humans than previously described., (© 2022 The Authors.)

Chimpanzees (Pan troglodytes) harbor rich assemblages of malaria parasites, including three species closely related to P. falciparum (sub-genus Laverania), the most malignant human malaria parasite. Here, we characterize the ecology and epidemiology of malaria infection in wild chimpanzee reservoirs. We used molecular assays to screen chimpanzee fecal samples, collected longitudinally and cross-sectionally from wild populations, for malaria parasite mitochondrial DNA. We found that chimpanzee malaria parasitism has an early age of onset and varies seasonally in prevalence. A subset of samples revealed Hepatocystis mitochondrial DNA, with phylogenetic analyses suggesting that Hepatocystis appears to cross species barriers more easily than Laverania. Longitudinal and cross-sectional sampling independently support the hypothesis that mean ambient temperature drives spatiotemporal variation in chimpanzee Laverania infection. Infection probability peaked at ~24.5 °C, consistent with the empirical transmission optimum of P. falciparum in humans. Forest cover was also positively correlated with spatial variation in Laverania prevalence, consistent with the observation that forest-dwelling Anophelines are the primary vectors. Extrapolating these relationships across equatorial Africa, we map spatiotemporal variation in the suitability of chimpanzee habitat for Laverania transmission, offering a hypothetical baseline indicator of human exposure risk., (© 2022. The Author(s).)

Understanding the affective lives of animals has been a long-standing challenge in science. Recent technological progress in infrared thermal imaging has enabled researchers to monitor animals' physiological states in real-time when exposed to ecologically relevant situations, such as feeding in the company of others. During social feeding, an individual's physiological states are likely to vary with the nature of the resource and perceptions of competition. Previous findings in chimpanzees have indicated that events perceived as competitive cause decreases in nasal temperatures, whereas the opposite was observed for cooperative interactions. Here, we tested how food resources and audience structure impacted on how social feeding events were perceived by wild chimpanzees. Overall, we found that nasal temperatures were lower when meat was consumed as compared to figs, consistent with the idea that social feeding on more contested resources is perceived as more dangerous and stressful. Nasal temperatures were significant affected by interactions between food type and audience composition, in particular the number of males, their dominance status, and their social bond status relative to the subject, while no effects for the presence of females were observed. Our findings suggest that male chimpanzees closely monitor and assess their social environment during competitive situations, and that infrared imaging provides an important complement to access psychological processes beyond observable social behaviours. This article is part of the theme issue 'Cognition, communication and social bonds in primates'.

The Interaction Engine Hypothesis postulates that humans have a unique ability and motivation for social interaction. A crucial juncture in the ontogeny of the interaction engine could be around 2-4 years of age, but observational studies of children in natural contexts are limited. These data appear critical also for comparison with non-human primates. Here, we report on focal observations on 31 children aged 2- and 4-years old in four preschools (10 h per child). Children interact with a wide range of partners, many infrequently, but with one or two close friends. Four-year olds engage in cooperative social interactions more often than 2-year olds and fight less than 2-year olds. Conversations and playing with objects are the most frequent social interaction types in both age groups. Children engage in social interactions with peers frequently (on average 13 distinct social interactions per hour) and briefly (28 s on average) and shorter than those of great apes in comparable studies. Their social interactions feature entry and exit phases about two-thirds of the time, less frequently than great apes. The results support the Interaction Engine Hypothesis, as young children manifest a remarkable motivation and ability for fast-paced interactions with multiple partners. This article is part of the theme issue 'Revisiting the human 'interaction engine': comparative approaches to social action coordination'.

Joint commitment, the feeling of mutual obligation binding participants in a joint action, is typically conceptualized as arising by the expression and acceptance of a promise. This account limits the possibilities of investigating fledgling forms of joint commitment in actors linguistically less well-endowed than adult humans. The feeling of mutual obligation is one aspect of joint commitment (the product ), which emerges from a process of signal exchange. It is gradual rather than binary; feelings of mutual obligation can vary in strength according to how explicit commitments are perceived to be. Joint commitment processes are more complex than simple promising, in at least three ways. They are affected by prior joint actions, which create precedents and conventions that can be embodied in material arrangements of institutions. Joint commitment processes also arise as solutions to generic coordination problems related to opening up, maintaining and closing down joint actions. Finally, during joint actions, additional, specific commitments are made piecemeal. These stack up over time and persist, making it difficult for participants to disengage from joint actions. These complexifications open up new perspectives for assessing joint commitment across species. This article is part of the theme issue 'Revisiting the human 'interaction engine': comparative approaches to social action coordination'.