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Wild chimpanzees drum on tree buttresses during dominance displays and travel, generating low-frequency sounds that are audible over distances of more than 1 km. Western chimpanzees (Pan troglodytes verus) in the Nimba Mountains of Guinea selectively choose trees and buttresses when drumming, potentially based on their resonant properties, suggesting that these chimpanzees are optimizing their drumming signals. We investigated whether male eastern chimpanzees (Pan troglodytes schweinfurthii) from the Waibira community in the Budongo Forest, Uganda, also show preferences in tree and buttress choice, exploring whether selectivity is a species-wide feature. We tested chimpanzee preferences for tree species and diameter, number of buttresses, and buttress area and width, by comparing trees and buttresses used in drumming bouts with nearby unused trees and buttresses. Waibira chimpanzees drummed preferentially on two tree species: the tropical hardwood Cynometra alexandrii and the softwood Chrysophyllum albidum. Chimpanzees selected trees with a larger diameter over nearby trees with a smaller diameter, and buttresses were more likely to be used for drumming if they had a larger area or larger width. These results suggest that chimpanzees in the Waibira community select trees and buttresses based on physical properties, most likely related to acoustically relevant characteristics. These findings support the argument that buttress drumming is a goal-directed behavior and contributes to our understanding of chimpanzees' use and optimization of their long-distance acoustic signals., (© 2024 The Author(s). American Journal of Primatology published by Wiley Periodicals LLC.)

Primates, the most colorful mammalian radiation, have previously served as an interesting model to test the functions and evolutionary drivers of variation in eye color. We assess the contribution of photo-regulatory and communicative functions to the external eye appearance of nine macaque species representing all the branches of their radiation. Macaques' well described social structure and wide geographical distribution make them interesting to explore. We find that (1) the posterior option of the anterior eyeball is more pigmented closer to the equator, suggesting photoprotective functions. We also find that (2) the temporal side of the eyeball is more heavily pigmented than the nasal side. This suggests that eyeball pigmentation in macaques is distributed to reduce damage to the corneal limbus. The inclusion of a translocated population of M. fuscata in our analyses also suggests that external eye appearance may change quickly, perhaps owing to phenotypic plasticity. We find no evidence that communicative functions drive variation in external eye appearance in macaques. These results suggest that the amount of light in a species' environment drives variation in eye coloration across macaque species. Furthermore, the geographical distribution of macaques hints at important factors that have yet to be accounted for, such as the reflectivity of the terrain a given species inhabits., Competing Interests: Declarations. Competing interests: The authors declare no competing interests., (© 2024. The Author(s).)

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).)

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.)

Social norms - rules governing which behaviours are deemed appropriate or inappropriate within a given community - are typically taken to be uniquely human. Recently, this position has been challenged by a number of philosophers, cognitive scientists, and ethologists, who have suggested that social norms may also be found in certain non-human animal communities. Such claims have elicited considerable scepticism from norm cognition researchers, who doubt that any non-human animals possess the psychological capacities necessary for normative cognition. However, there is little agreement among these researchers about what these psychological prerequisites are. This makes empirical study of animal social norms difficult, since it is not clear what we are looking for and thus what should count as behavioural evidence for the presence (or absence) of social norms in animals. To break this impasse, we offer an approach that moves beyond contested psychological criteria for social norms. This approach is inspired by the animal culture research program, which has made a similar shift away from heavily psychological definitions of 'culture' to become organised around a cluster of more empirically tractable concepts of culture. Here, we propose an analogous set of constructs built around the core notion of a normative regularity, which we define as a socially maintained pattern of behavioural conformity within a community. We suggest methods for studying potential normative regularities in wild and captive primates. We also discuss the broader scientific and philosophical implications of this research program with respect to questions of human uniqueness, animal welfare and conservation., (© 2024 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

Many species communicate by combining signals into multimodal combinations. Elephants live in multi-level societies where individuals regularly separate and reunite. Upon reunion, elephants often engage in elaborate greeting rituals, where they use vocalisations and body acts produced with different body parts and of various sensory modalities (e.g., audible, tactile). However, whether these body acts represent communicative gestures and whether elephants combine vocalisations and gestures during greeting is still unknown. Here we use separation-reunion events to explore the greeting behaviour of semi-captive elephants (Loxodonta africana). We investigate whether elephants use silent-visual, audible, and tactile gestures directing them at their audience based on their state of visual attention and how they combine these gestures with vocalisations during greeting. We show that elephants select gesture modality appropriately according to their audience's visual attention, suggesting evidence of first-order intentional communicative use. We further show that elephants integrate vocalisations and gestures into different combinations and orders. The most frequent combination consists of rumble vocalisations with ear-flapping gestures, used most often between females. By showing that a species evolutionarily distant to our own primate lineage shows sensitivity to their audience's visual attention in their gesturing and combines gestures with vocalisations, our study advances our understanding of the emergence of first-order intentionality and multimodal communication across taxa., (© 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.)

Current methodologies present significant hurdles to understanding patterns in the gestural communication of individuals, populations, and species. To address this issue, we present a bottom-up data collection framework for the study of gesture: GesturalOrigins. By "bottom-up", we mean that we minimise a priori structural choices, allowing researchers to define larger concepts (such as 'gesture types', 'response latencies', or 'gesture sequences') flexibly once coding is complete. Data can easily be re-organised to provide replication of, and comparison with, a wide range of datasets in published and planned analyses. We present packages, templates, and instructions for the complete data collection and coding process. We illustrate the flexibility that our methodological tool offers with worked examples of (great ape) gestural communication, demonstrating differences in the duration of action phases across distinct gesture action types and showing how species variation in the latency to respond to gestural requests may be revealed or masked by methodological choices. While GesturalOrigins is built from an ape-centred perspective, the basic framework can be adapted across a range of species and potentially to other communication systems. By making our gesture coding methods transparent and open access, we hope to enable a more direct comparison of findings across research groups, improve collaborations, and advance the field to tackle some of the long-standing questions in comparative gesture research., (© 2023. The Author(s).)

We report the presence of habitual ground nesting in a newly studied East African chimpanzee (Pan troglodytes schweinfurthii) population in the Bugoma Central Forest Reserve, Uganda. Across a 2-year period, we encountered 891 night nests, 189 of which were classified as ground nests, a rate of ~21%. We find no preliminary evidence of socio-ecological factors that would promote its use and highlight local factors, such as high incidence of forest disturbance due to poaching and logging, which appear to make its use disadvantageous. While further study is required to establish whether this behavior meets the strict criteria for nonhuman animal culture, we support the argument that the wider use of population and group-specific behavioral repertoires in flagship species, such as chimpanzees, offers a tool to promote the urgent conservation action needed to protect threatened ecosystems, including the Bugoma forest., (© 2023 The Authors. American Journal of Primatology published by Wiley Periodicals LLC.)

Postings on social media on Twitter (now X), BioAnthropology News (Facebook), and other venues, as well as recent publications in prominent journals, show that primatologists, ecologists, and other researchers are questioning the terms "Old World" and "New World" due to their colonial implications and history. The terms are offensive if they result in erasing Indigenous voices and history, ignoring the fact that Indigenous peoples were in the Americas long before European colonization. Language use is not without context, but alternative terminology is not always obvious and available. In this perspective, we share opinions expressed by an international group of primatologists who considered questions about the use of these terms, whether primatologists should adjust language use, and how to move forward. The diversity of opinions provides insight into how conventional terms used in primatological research and conservation may impact our effectiveness in these domains., (© 2023. The Author(s).)

In the hall of animal oddities, the reindeer ( Rangifer tarandus ) is the only mammal with a color-shifting tapetum lucidum and the only ruminant with a lichen-dominated diet. These puzzling traits coexist with yet another enigma--ocular media that transmit up to 60% of ultraviolet (UV) light, enough to excite the cones responsible for color vision. It is unclear why any day-active circum-Arctic mammal would benefit from UV visual sensitivity, but it could improve detection of UV-absorbing lichens against a background of UV-reflecting snows, especially during the extended twilight hours of winter. To explore this idea and advance our understanding of reindeer visual ecology, we recorded the reflectance spectra of several ground-growing (terricolous), shrubby (fruticose) lichens in the diets of reindeer living in Cairngorms National Park, Scotland., Competing Interests: The authors declared no potential conflicts of interest with respect to the research, authorship, and/or publication of this article., (© The Author(s) 2023.)

Chimpanzees were once thought to sleep primarily in the trees, but recent studies indicate that some populations also construct terrestrial night nests. This behavior has relevance not only to understanding the behavioral diversity of Pan troglodytes, but also to the conservation of the species, given that nest encounter rates are often used to estimate great ape population densities. A proper estimate of decay rates for ground nests is necessary for converting the encounter rate of nests to the density of weaned chimpanzees. Here we present the results of the first systematic comparative study between the decay rates of arboreal and terrestrial chimpanzee nests, from the Bugoma Central Forest Reserve in western Uganda. We followed the decay of 56 ground and 51 tree nests in eight nest groups between April 2020 and October 2021. For 15 of the ground and 19 of the tree nests, we collected detailed information on the condition of the nests every two weeks; we checked the remaining 73 nests only twice. On average, ground nests lasted 238 days versus 276 days for tree nests (p = 0.05). Of the 107 total nests surveyed, 51% of tree and 64% of ground nests had disappeared after six months. Based on our results, we propose a modification of the formula used to convert nest density into chimpanzee density. Our results highlight the importance of taking into account potential differences in decay rates between ground versus tree nests, which will likely influence our understanding of the distribution of ground nesting behavior in chimpanzee across tropical Africa, as well as our estimations of the densities of ground nesting populations., (© 2023 The Authors. American Journal of Primatology published by Wiley Periodicals LLC.)

Object interactions play an important role in human communication but the extent to which nonhuman primates incorporate objects in their social interactions remains unknown. To better understand the evolution of object use, this study explored how objects are used in social interactions in semi-wild chimpanzees (Pan troglodytes). We used an observational approach focusing on naturally occurring object actions where we examined their use and tested whether the production of object actions was influenced by the recipients' visual attention as well as by colony membership. The results show that chimpanzees adjusted both the type of object used, and the modality of object actions to match the visual attention of the recipient, as well as colony differences in the use of targeted object actions. These results provide empirical evidence highlighting that chimpanzees use objects in diverse ways to communicate with conspecifics and that their use may be shaped by social factors, contributing to our understanding of the evolution of human nonverbal communication, language, and tool use., (© 2023. The Author(s).)

Studying animal behaviour allows us to understand how different species and individuals navigate their physical and social worlds. Video coding of behaviour is considered a gold standard: allowing researchers to extract rich nuanced behavioural datasets, validate their reliability, and for research to be replicated. However, in practice, videos are only useful if data can be efficiently extracted. Manually locating relevant footage in 10,000 s of hours is extremely time-consuming, as is the manual coding of animal behaviour, which requires extensive training to achieve reliability. Machine learning approaches are used to automate the recognition of patterns within data, considerably reducing the time taken to extract data and improving reliability. However, tracking visual information to recognise nuanced behaviour is a challenging problem and, to date, the tracking and pose-estimation tools used to detect behaviour are typically applied where the visual environment is highly controlled. Animal behaviour researchers are interested in applying these tools to the study of wild animals, but it is not clear to what extent doing so is currently possible, or which tools are most suited to particular problems. To address this gap in knowledge, we describe the new tools available in this rapidly evolving landscape, suggest guidance for tool selection, provide a worked demonstration of the use of machine learning to track movement in video data of wild apes, and make our base models available for use. We use a pose-estimation tool, DeepLabCut, to demonstrate successful training of two pilot models of an extremely challenging pose estimate and tracking problem: multi-animal wild forest-living chimpanzees and bonobos across behavioural contexts from hand-held video footage. With DeepWild we show that, without requiring specific expertise in machine learning, pose estimation and movement tracking of free-living wild primates in visually complex environments is an attainable goal for behavioural researchers., (© 2023 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Mechanisms of inheritance remain poorly defined for many fitness-mediating traits, especially in long-lived animals with protracted development. Using 6,123 urinary samples from 170 wild chimpanzees, we examined the contributions of genetics, non-genetic maternal effects, and shared community effects on variation in cortisol levels, an established predictor of survival in long-lived primates. Despite evidence for consistent individual variation in cortisol levels across years, between-group effects were more influential and made an overwhelming contribution to variation in this trait. Focusing on within-group variation, non-genetic maternal effects accounted for 8% of the individual differences in average cortisol levels, significantly more than that attributable to genetic factors, which was indistinguishable from zero. These maternal effects are consistent with a primary role of a shared environment in shaping physiology. For chimpanzees, and perhaps other species with long life histories, community and maternal effects appear more relevant than genetic inheritance in shaping key physiological traits., (© 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).)

Competing Interests: Declaration of Competing Interest The author declares that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

In the comparative study of human and nonhuman communication, ape gesturing provided the first demonstrations of flexible, intentional communication outside human language. Rich repertoires of these gestures have been described in all ape species, bar one: us. Given that the majority of great ape gestural signals are shared, and their form appears biologically inherited, this creates a conundrum: Where did the ape gestures go in human communication? Here, we test human recognition and understanding of 10 of the most frequently used ape gestures. We crowdsourced data from 5,656 participants through an online game, which required them to select the meaning of chimpanzee and bonobo gestures in 20 videos. We show that humans may retain an understanding of ape gestural communication (either directly inherited or part of more general cognition), across gesture types and gesture meanings, with information on communicative context providing only a marginal improvement in success. By assessing comprehension, rather than production, we accessed part of the great ape gestural repertoire for the first time in adult humans. Cognitive access to an ancestral system of gesture appears to have been retained after our divergence from other apes, drawing deep evolutionary continuity between their communication and our own., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2023 Graham, Hobaiter. 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.)

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).)

The last decades have seen major advances in the study of gestures both in humans and non-human primates. In this paper, we seriously examine the idea that there may be gestural form types that are shared across great ape species, including humans, which may underlie gestural universals, both in form and meaning. We focus on one case study, the hand fling gesture common to chimpanzees and humans, and provide a semantic analysis of this gesture., (© 2022 the author(s), published by De Gruyter, Berlin/Boston.)

Two language laws have been identified as consistent patterns shaping animal behaviour, both acting on the organizational level of communicative systems. Zipf's law of brevity describes a negative relationship between behavioural length and frequency. Menzerath's law defines a negative correlation between the number of behaviours in a sequence and average length of the behaviour composing it. Both laws have been linked with the information-theoretic principle of compression, which tends to minimize code length. We investigated their presence in a case study of male chimpanzee sexual solicitation gesture. We failed to find evidence supporting Zipf's law of brevity, but solicitation gestures followed Menzerath's law: longer sequences had shorter average gesture duration. Our results extend previous findings suggesting gesturing may be limited by individual energetic constraints. However, such patterns may only emerge in sufficiently large datasets. Chimpanzee gestural repertoires do not appear to manifest a consistent principle of compression previously described in many other close-range systems of communication. Importantly, the same signallers and signals were previously shown to adhere to these laws in subsets of the repertoire when used in play; highlighting that, in addition to selection on the signal repertoire, ape gestural expression appears shaped by factors in the immediate socio-ecological context., (© 2022 The Authors.)

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).

External eye appearance across primate species is diverse in shape and colouration, yet we still lack an explanation for the drivers of such diversity. Here we quantify substantial interspecific variation in eye shape and colouration across 77 primate species representing all extant genera of anthropoid primates. We reassess a series of hypotheses aiming to explain ocular variation in horizontal elongation and in colouration across species. Heavier body weight and terrestrial locomotion are associated with elongated eye outlines. Species living closer to the equator present more pigmented conjunctivae, suggesting photoprotective functions. Irises become bluer in species living further away from the equator, adding to existing literature supporting a circadian clock function for bluer irises. These results shift the current focus from communicative, to ecological factors in driving variation in external eye appearance in anthropoid primates. They also highlight the possibility that similar ecological factors contributed to selection for blue eyes in ancestral human populations living in northern latitudes., (© 2022. The Author(s).)

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.)

Opinion piece: ape gestures are made intentionally, inviting parallels with human language; but how similar are their gestures to words? Here we ask this in three ways, considering: flexibility and ambiguity, first- and second-order intentionality, and usage in interactive exchanges. Many gestures are used to achieve several, often very distinct, goals. Such apparent ambiguity in meaning is potentially disruptive for communication, but-as with human language-situational and interpersonal context may largely resolve the intended meaning. Our evidence for first-order intentional use of gesture is abundant, but how might we establish a case for the second-order intentional use critical to language? Finally, words are rarely used in tidy signal-response sequences but are exchanged in back-and-forth interaction. Do gestures share this property? In this paper, we examine these questions and set out ways in which they can be resolved, incorporating data from wild chimpanzees. This article is part of the theme issue 'Cognition, communication and social bonds in primates'.

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'.

In human fission-fusion societies, ritualized non-linguistic signal exchanges that include gestures, vocalizations, and facial expressions are regularly observed at both arrivals (greetings) and departures (leave-takings). These communicative events play an important role in the formation and maintenance of social relationships. Wild chimpanzees also form large communities that split into smaller fluid parties during daily activities, with individuals moving freely between them. However, in chimpanzees only greetings have been reported. This study explores signal exchanges in the Bossou chimpanzee community during fissions (departures) and fusions (arrivals) given an individual's social rank, kinship, position as traveller or party-member, the level of potential threat, and the party size and presence of mature males. We analysed three time periods (1993-1994; 2003-2004; 2013-2014) during which the composition and social hierarchy of the community varied. We show that the occurrence and form of communication during fission and fusion events are mediated by social factors, including rank, kinship, and party size and composition. Individuals were more likely to communicate during fusions than during fissions, communication was more likely to be produced towards a higher-ranking individual and to non-kin individuals, but the tendency to communicate in general increased with an increase in social rank. The presence of more individuals, and in particular mature males, decreased the likelihood of communication. Communication during fusions supported patterns reported in previous studies on greetings, and our results support the argument that, if present, leave-takings are not a common feature of chimpanzee social interactions. Current methodological difficulties regarding the function of declarative signals hinder our ability to discriminate potential parting rituals within communication before departures. Given similar methodological difficulties, we also provide a note of caution in the interpretation of all signals produced during fusions as 'greetings'., (© 2021. Japan Monkey Centre.)

It has been suggested that non-human primates can respond to deceased conspecifics in ways that suggest they experience psychological states not unlike humans, some of which could indicate they exhibit a notion of death. Here, we report long-term demographic data from two East African chimpanzee groups. During a combined 40-year observation period, we recorded 191 births of which 68 died in infancy, mostly within the first year. We documented the post-mortem behaviour of the mothers and describe nine occasions where Budongo chimpanzee mothers carried infants for 1-3 days after their death, usually until the body started to decompose. We also observed three additional cases of extended carrying lasting for more than 2 weeks, one of which was followed by the unusual extended carrying of an object and another which lasted 3 months. In each case, the corpses mummified. In addition, we report four instances of recurring dead-infant carrying by mothers, three of whom carried the corpse for longer during the second instance. We discuss these observations in view of functional hypotheses of dead-infant carrying in primates and the potential proximate mechanisms involved in this behaviour., (© 2022. The Author(s).)

Access to resources shapes species' physiology and behaviour. Water is not typically considered a limiting resource for rainforest-living chimpanzees; however, several savannah and savannah-woodland communities show behavioural adaptations to limited water. Here, we provide a first report of habitual well-digging in a rainforest-living group of East African chimpanzees (Pan troglodytes schweinfurthii) and suggest that it may have been imported into the community's behavioural repertoire by an immigrant female. We describe the presence and frequency of well-digging and related behaviour, and suggest that its subsequent spread in the group may have involved some degree of social learning. We highlight that subsurface water is a concealed resource, and that the limited spread of well-digging in the group may highlight the cognitive, rather than physical, challenges it presents in a rainforest environment., (© 2022. The Author(s).)

Albinism-the congenital absence of pigmentation-is a very rare phenomenon in animals due to the significant costs to fitness of this condition. Both humans and non-human individuals with albinism face a number of challenges, such as reduced vision, increased exposure to ultraviolet radiation, or compromised crypticity resulting in an elevated vulnerability to predation. However, while observations of social interactions involving individuals with albinism have been observed in wild non-primate animals, such interactions have not been described in detail in non-human primates (hereafter, primates). Here, we report, to our knowledge, the first sighting of an infant with albinism in wild chimpanzees (Pan troglodytes schweinfurthii), including social interactions between the infant, its mother, and group members. We also describe the subsequent killing of the infant by conspecifics as well as their behavior towards the corpse following the infanticide. Finally, we discuss our observations in relation to our understanding of chimpanzee behavior or attitudes towards individuals with very conspicuous appearances., (© 2021 The Authors. American Journal of Primatology published by Wiley Periodicals LLC.)

Over the last 30 years, most research on non-human primate gestural communication has been produced by psychologists, which has shaped the questions asked and the methods used. These researchers have drawn on concepts from philosophy, linguistics, anthropology, and ethology, but despite these broad influences the field has neglected to situate gestures into the socio-ecological context in which the diverse species, individuals, and social-units exist. In this review, we present current knowledge about great ape gestural communication in terms of repertoires, meanings, and development. We fold this into a conversation about variation in other types of ape social behaviour to identify areas for future research on variation in gestural communication. Given the large variation in socio-ecological factors across species and social-units (and the individuals within these groups), we may expect to find different preferences for specific gesture types; different needs for communicating specific meanings; and different rates of encountering specific contexts. New tools, such as machine-learning based automated movement tracking, may allow us to uncover potential variation in the speed and form of gesture actions or parts of gesture actions. New multi-group multi-generational datasets provide the opportunity to apply analyses, such as Bayesian modelling, which allows us to examine these rich behavioural landscapes. Together, by expanding our questions and our methods, researchers may finally be able to study great ape gestures from the perspective of the apes themselves and explore what this gestural communication system reveals about apes' thinking and experience of their world., Competing Interests: No potential conflict of interest was reported by the authors., (© 2022 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group.)

Gestural communication permeates all domains of chimpanzees' social life and is intentional in use. However, we still have only limited information on how young apes develop the sociocognitive skills needed for intentional communication. In this cross-sectional study, we document the development of behavioral adjustment to the recipient's visual attention-considered a hallmark of intentional communication-in wild immature chimpanzees' gestural communication. We studied 11 immature chimpanzees (Pan troglodytes schweinfurthii): three infants, four juveniles, and four adolescents gesturing towards their mother. We quantified silent-visual, audible, and contact gestures indexed to maternal visual attention and inattention. We investigated unimodal adjustment, defined by the capacity of young chimpanzees to deploy fewer silent-visual signals when their mothers did not show full visual attention towards them as compared with when they did. We then examined cross-modal adjustment, defined as the capacity of chimpanzees to deploy more audible-or-contact gestures than silent-visual gestures in the condition where their mothers did not show full visual attention as compared to when they did. Our results show a gradual decline in the use of silent-visual gestures when the mother is not visually attentive with increasing age. The absence of silent-visual gesture production toward a visually inattentive recipient (complete unimodal adjustment) was not fully in place until adolescence. Immature chimpanzees used more audible-or-contact gestures than silent-visual ones when their mothers did not show visual attention and vice-versa when they did. This cross-modal adjustment was expressed in juveniles and adolescents but not in infants. Overall, this study shows that infant chimpanzees were limited in their sensitivity to maternal attention when gesturing, whereas adolescent chimpanzees adjusted their communication appropriately. Juveniles present an intermediate pattern with cross-modal adjustment preceding unimodal adjustment and with variability in the age of onset., (© 2021 Wiley Periodicals LLC.)

The global lockdown to mitigate COVID-19 pandemic health risks has altered human interactions with nature. Here, we report immediate impacts of changes in human activities on wildlife and environmental threats during the early lockdown months of 2020, based on 877 qualitative reports and 332 quantitative assessments from 89 different studies. Hundreds of reports of unusual species observations from around the world suggest that animals quickly responded to the reductions in human presence. However, negative effects of lockdown on conservation also emerged, as confinement resulted in some park officials being unable to perform conservation, restoration and enforcement tasks, resulting in local increases in illegal activities such as hunting. Overall, there is a complex mixture of positive and negative effects of the pandemic lockdown on nature, all of which have the potential to lead to cascading responses which in turn impact wildlife and nature conservation. While the net effect of the lockdown will need to be assessed over years as data becomes available and persistent effects emerge, immediate responses were detected across the world. Thus, initial qualitative and quantitative data arising from this serendipitous global quasi-experimental perturbation highlights the dual role that humans play in threatening and protecting species and ecosystems. Pathways to favorably tilt this delicate balance include reducing impacts and increasing conservation effectiveness., Competing Interests: Authors declare no competing interests., (© 2021 Published by Elsevier Ltd.)

Catherine Hobaiter, John Walter Akankwasa, Geresomu Muhumuza, Moreen Uwimbabazi and Inza Koné discuss the importance of local specialists in science., (Copyright © 2021 Elsevier Inc. All rights reserved.)

Comparative psychology provides important contributions to our understanding of the origins of human language. The presence of common features in human and nonhuman primate communication can be used to suggest the evolutionary trajectories of potential precursors to language. However, to do so effectively, our findings must be comparable across diverse species. This systematic review describes the current landscape of data available from studies of gestural communication in human and nonhuman primates that make an explicit connection to language evolution. We found a similar number of studies on human and nonhuman primates, but that very few studies included data from more than one species. As a result, evolutionary inferences remain restricted to comparison across studies. We identify areas of focus, bias, and apparent gaps within the field. Different domains have been studied in human and nonhuman primates, with relatively few nonhuman primate studies of ontogeny and relatively few human studies of gesture form. Diversity in focus, methods, and socio-ecological context fill important gaps and provide nuanced understanding, but only where the source of any difference between studies is transparent. Many studies provide some definition for their use of gesture; but definitions of gesture, and in particular, criteria for intentional use, are absent in the majority of human studies. We find systematic differences between human and nonhuman primate studies in the research scope, incorporation of other modalities, research setting, and study design. We highlight eight particular areas in a call to action through which we can strengthen our ability to investigate gestural communication's contribution within the evolutionary roots of human language., (© 2021 Wiley Periodicals LLC.)

Is it possible to slow the rate of ageing, or do biological constraints limit its plasticity? We test the 'invariant rate of ageing' hypothesis, which posits that the rate of ageing is relatively fixed within species, with a collection of 39 human and nonhuman primate datasets across seven genera. We first recapitulate, in nonhuman primates, the highly regular relationship between life expectancy and lifespan equality seen in humans. We next demonstrate that variation in the rate of ageing within genera is orders of magnitude smaller than variation in pre-adult and age-independent mortality. Finally, we demonstrate that changes in the rate of ageing, but not other mortality parameters, produce striking, species-atypical changes in mortality patterns. Our results support the invariant rate of ageing hypothesis, implying biological constraints on how much the human rate of ageing can be slowed.

Compared with most mammals, postnatal development in great apes is protracted, presenting both an extended period of phenotypic plasticity to environmental conditions and the potential for sustained mother-offspring and/or sibling conflict over resources. Comparisons of cortisol levels during ontogeny can reveal physiological plasticity to species or population specific socioecological factors and in turn how these factors might ameliorate or exaggerate mother-offspring and sibling conflict. Here, we examine developmental patterns of cortisol levels in two wild chimpanzee populations (Budongo and Taï), with two and three communities each, and one wild bonobo population (LuiKotale), with two communities. Both species have similar juvenile life histories. Nonetheless, we predicted that key differences in socioecological factors, such as feeding competition, would lead to interspecific variation in mother-offspring and sibling conflict and thus variation in ontogenetic cortisol patterns. We measured urinary cortisol levels in 1394 samples collected from 37 bonobos and 100 chimpanzees aged up to 12 years. The significant differences in age-related variation in cortisol levels appeared population specific rather than species specific. Both bonobos and Taï chimpanzees had comparatively stable and gradually increasing cortisol levels throughout development; Budongo chimpanzees experienced declining cortisol levels before increases in later ontogeny. These age-related population differences in cortisol patterns were not explained by mother-offspring or sibling conflict specifically; instead, the comparatively stable cortisol patterns of bonobos and Taï chimpanzees likely reflect a consistency in experience of competition and the social environment compared with Budongo chimpanzees, where mothers may adopt more variable strategies related to infanticide risk and resource availability. The clear population-level differences within chimpanzees highlight potential intraspecific flexibility in developmental processes in apes, suggesting the flexibility and diversity in rearing strategies seen in humans may have a deep evolutionary history., Competing Interests: Competing interests None of the authors have competing interests to report., (Copyright © 2020 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Interview with Cat Hobaiter, who studies the evolution of communication, cognition and social behaviour in apes at the University of St Andrews and through field studies in the Budongo Forest in Uganda., (Copyright © 2020.)

Speech is a human hallmark, but its evolutionary origins continue to defy scientific explanation. Recently, the open-close mouth rhythm of 2-7 Hz (cycles/second) characteristic of all spoken languages has been identified in the orofacial signals of several nonhuman primate genera, including orangutans, but evidence from any of the African apes remained missing. Evolutionary continuity for the emergence of speech is, thus, still inconclusive. To address this empirical gap, we investigated the rhythm of chimpanzee lip-smacks across four populations (two captive and two wild). We found that lip-smacks exhibit a speech-like rhythm at approximately 4 Hz, closing a gap in the evidence for the evolution of speech-rhythm within the primate order. We observed sizeable rhythmic variation within and between chimpanzee populations, with differences of over 2 Hz at each level. This variation did not result, however, in systematic group differences within our sample. To further explore the phylogenetic and evolutionary perspective on this variability, inter-individual and inter-population analyses will be necessary across primate species producing mouth signals at speech-like rhythm. Our findings support the hypothesis that speech recruited ancient primate rhythmic signals and suggest that multi-site studies may still reveal new windows of understanding about these signals' use and production along the evolutionary timeline of speech.

Maternal cannibalism has been reported in several animal taxa, prompting speculations that the behavior may be part of an evolved strategy. In chimpanzees, however, maternal cannibalism has been conspicuously absent, despite high levels of infant mortality and reports of non-maternal cannibalism. The typical response of chimpanzee mothers is to abandon their deceased infant, sometimes after prolonged periods of carrying and grooming the corpse. Here, we report two anomalous observations of maternal cannibalism in communities of wild chimpanzees in Uganda and Ivory Coast and discuss the evolutionary implications. Both infants likely died under different circumstances; one apparently as a result of premature birth, the other possibly as a result of infanticide. In both cases, the mothers consumed parts of the corpse and participated in meat sharing with other group members. Neither female presented any apparent signs of ill health before or after the events. We concluded that, in both cases, cannibalizing the infant was unlikely due to health-related issues by the mothers. We discuss these observations against a background of chimpanzee mothers consistently refraining from maternal cannibalism, despite ample opportunities and nutritional advantages. We conclude that maternal cannibalism is extremely rare in this primate, likely due to early and strong mother-offspring bond formation, which may have been profoundly disrupted in the current cases.