Over six decades of research on wild baboons and their close relatives (collectively, the African papionins) have uncovered substantial variation in their behavior and social systems. While most papionins form discrete social groups (single-level societies), a few others form small social units that are nested within larger supergroups (multi-level societies). These two systems are generally thought to be qualitatively distinct, but data from wild populations increasingly suggest that there may be areas of overlap. To quantify this potential gradient in social structure, a more systematic, comparative analysis is needed. Here, we constructed a database of behavioral and demographic records spanning 135 group-years, 28 social groups, 13 long-term field studies, and 11 species to quantify variation in grooming network structure and identify the individual and dyadic properties (e.g., kinship and social status effects) that underlie this variation. Consistent with accumulating field observations, the single-level species could be divided into two categories: cohesive and cliquish. Cohesive single-level networks were dense, kin-biased, and moderately rank-structured, while cliquish single-level networks were more differentiated, slightly more kin-biased, and strongly rank-structured. As expected, multi-level networks were very modular and shaped by females' ties to specific dominant males but varied in their kin biases. Taken together, these data suggest that in the African papionins i) kin and rank biases are widespread but vary in their strength; ii) male-centered subgroups are exclusive to multi-level systems; and iii) increases in network modularity can emerge in response to heightened nepotism and male-centered clustering.

There is a vast and ever-accumulating amount of behavioural data on individually recognised animals, an incredible resource to shed light on the ecological and evolutionary drivers of variation in animal behaviour. Yet, the full potential of such data lies in comparative research across taxa with distinct life histories and ecologies. Substantial challenges impede systematic comparisons, one of which is the lack of persistent, accessible and standardised databases. Big-team approaches to building standardised databases offer a solution to facilitating reliable cross-species comparisons. By sharing both data and expertise among researchers, these approaches ensure that valuable data, which might otherwise go unused, become easier to discover, repurpose and synthesise. Additionally, such large-scale collaborations promote a culture of sharing within the research community, incentivising researchers to contribute their data by ensuring their interests are considered through clear sharing guidelines. Active communication with the data contributors during the standardisation process also helps avoid misinterpretation of the data, ultimately improving the reliability of comparative databases. Here, we introduce MacaqueNet, a global collaboration of over 100 researchers (https://macaquenet.github.io/) aimed at unlocking the wealth of cross-species data for research on macaque social behaviour. The MacaqueNet database encompasses data from 1981 to the present on 61 populations across 14 species and is the first publicly searchable and standardised database on affiliative and agonistic animal social behaviour. We describe the establishment of MacaqueNet, from the steps we took to start a large-scale collective, to the creation of a cross-species collaborative database and the implementation of data entry and retrieval protocols. We share MacaqueNet's component resources: an R package for data standardisation, website code, the relational database structure, a glossary and data sharing terms of use. With all these components openly accessible, MacaqueNet can act as a fully replicable template for future endeavours establishing large-scale collaborative comparative databases.

How primates sense, cognize, and respond to death, and are affected by it, are broad research themes within primate thanatology. When primate thanatology adopts a comparative framework, it can illuminate the diversity and evolutionary continuity of psycho-neuro-emotional responses to death and dying across primate species, including humans. Evolutionary or comparative thanatology has recently gained momentum, but it lacks methodological consistency, quantitative rigor, and coherent reporting standards. We attempt to address the lack of standardized approaches by introducing an individual-level comparative framework and employing standard ethological methods, accompanied by well-defined behavioral variables. We argue that the proposed framework can capture individual-specific responses to conspecific death and enable hypothesis-driven studies. The framework is essential to examine the large number of hypotheses that have accumulated in the field.

In animal social groups, the extent to which individuals consistently win agonistic interactions and their ability to monopolize resources represent 2 core aspects of their competitive regime. However, whether these two aspects are closely correlated within groups has rarely been studied. Here, we tested the hypothesis that hierarchy steepness, which is generally used to represent power differentials between group members, predicts the variation in the distribution of fitness-related benefits (i.e. fecundity, infant survival, mating success, and feeding success) in relation to individual dominance ranks. We tested this hypothesis in primate groups using comparative phylogenetic meta-analytical techniques. Specifically, we reviewed published and unpublished studies to extract data on individual dominance ranks, their access to fitness-related benefits, and hierarchy steepness. We collected and included in our analysis a total of 153 data points, representing 27 species (including 2 chimpanzee sub-species). From these, we used 4 common methods to measure individual dominance ranks and hierarchy steepness, i.e. Dij-based normalized David's scores, randomized Elo-ratings, and David's scores and Elo-ratings estimated in Bayesian frameworks. We found that hierarchy steepness had no effect on the strength of the relationship between dominance rank and access to fitness-related benefits. Our results suggest that hierarchy steepness does not reflect between-group variation in the extent to which individual dominance affects the acquisition of fitness-related benefits in primates. Although the ability to win agonistic encounters is essential, we speculate that other behavioral strategies adopted by individuals may play crucial roles in resource acquisition in animal competitive regimes. In social animals, group members compete to attain dominant positions. Dominant individuals are expected to have better access to key resources, like food or mating. In our study, we show that the strength of the dominance hierarchy does not affect the distribution of key resources in primates. The distribution of social bonds and agonistic support between group members may weaken the effect of dominance hierarchy on how key resources are accessed by dominants and subordinates.

Abstract The purpose of our research was to identify the primary personality traits of bonnet macaques using an observer-based questionnaire and to assess the accuracy of this method in predicting behavior. Additionally, we examined the relationship between personality traits and variables such as sex, age, and rank. We rated 25 males and 38 females on 50 personality questionnaire items using a 7-point scale, with reliable ratings and good agreement among raters (the lowest of the calculated ICC (3:k) index ≥ 0.62). Behavioral observations of 38 females over two-years were analyzed. Exploratory factor analysis revealed five traits: Friendliness, Aggression, Playfulness, Fearfulness, and Opportunism. Every trait correlated with a certain number of observed behaviors. Our results suggest that the questionnaire-based method is sufficiently valid as it confirmed high behavioral convergence with Fearfulness, Playfulness and Aggressiveness. Further studies are needed to understand the functional value of behaviors associated with Friendliness and Opportunism. Our study confirmed a tolerant social style of bonnet macaques, characterized by similar Aggression levels in both males and females, and lack of difference in Aggression level between subadult and adult males. Moreover, we identified a species-characteristic trait Opportunism which, unlike in other studied macaques, did not involve aggression-related characteristics.

Comparative studies reliant on single personality surveys to rate wild primates are scarce yet remain critical for developing a holistic comparative understanding of personality. Differences in survey design, item exclusion, and factor selection impede cross-study comparisons. To address these challenges, we used consistently collected data to assess personality trait structures in wild rhesus (Macaca mulatta), bonnet (M. radiata), and long-tailed (M. fascicularis) macaques that varied in their degree of phylogenetic closeness, species-typical social styles, and anthropogenic exposure in urban or urban-rural environments. We administered 51-item personality surveys to familiar raters, and, after reliability and structure screenings, isolated 4-5 factor solutions among the species. Four consistent factors emerged: Confident, Sociable, Active, and Irritable/Equable. This latter factor had differential expression across species. Item composition of the Irritable/Equable factor was consistent with their anticipated differences in social styles, but confounded by cross-site anthropogenic variation. We also administered a 43-item survey confined to human-primate situations which paralleled our findings of social style variation, while also exhibiting variation that aligned with population differences in human density. Our findings indicate that macaque personality trait structures may be emergent outcomes of evolutionary and/or socioecological processes, but further research is needed to parse these processes' relative contributions.

Mother–offspring bonds in primates often last for life and shape the behavior of both mother and offspring. Therefore, the death of an infant may trigger an emotional response from its mother akin to human grief, which is characterized by active distress and passive depression. Our goal was to investigate whether the behavior of female bonnet macaques ( Macaca radiata ) changed after the loss of their infants. In the Thenmala area, southern India, between 2020 and 2022, we observed 18 mothers in two groups who lost, in total, 19 infants and three juveniles aged < 18 months. We compared affiliative, aggressive, and stress-related behavior of females before and after the death of their infants. We also examined individual variation in responses to infant loss. After the death of their infants, females withdrew socially, stayed on the periphery of the group, and had fewer companions. Bereaved females were less involved in grooming and hugging and initiated aggressive behavior more frequently than before they lost their infants. Moreover, females who lost infants showed frequent stress-related behaviors, such as self-scratching and yawning compared with when their infants were alive. Our results suggest that female bonnet macaques react to deaths of their infants in a manner that is consistent with the interpretation of grief in humans and other primates.

Though uncommon, adoption of orphaned infants has been observed in both wild and captive non-human primates. In two groups of wild bonnet macaques ( Macaca radiata ), we observed five instances of infants being cared for after they lost their mothers at a pre-weaning age (< 6 months). Orphaned infants had one or more caregivers (juvenile, subadult, and adult female or male) involved in carrying, grooming, hugging, and protecting them. Adoption did not appear to be related to the age/sex class of the infant, or directly to the mother’s rank. Although the dominance rank of the mother of an orphaned infant did not have a direct effect on orphan survivorship, it determined the number of caregivers available to the orphaned infant, and infant survivorship was positively related to the number of caregivers of the orphaned infant. Thus, survivorship was likely a function of the mother’s sociality. Two other infants born to high-ranking mothers were also adopted by more individuals and survived longer than the infants of low-ranking mothers.

Pandemics caused by pathogens that originate in wildlife highlight the importance of understanding the behavioral ecology of disease outbreaks at human-wildlife interfaces. Specifically, the relative effects of human-wildlife and wildlife-wildlife interactions on disease outbreaks among wildlife populations in urban and peri-urban environments remain unclear. We used social network analysis and epidemiological Susceptible-Infected-Recovered models to simulate zooanthroponotic outbreaks, through wild animals' joint propensities to co-interact with humans, and their social grooming of conspecifics. On 10 groups of macaques (Macaca spp.) in peri-urban environments in Asia, we collected behavioral data using event sampling of human-macaque interactions within the same time and space, and focal sampling of macaques' social interactions with conspecifics and overall anthropogenic exposure. Model-predicted outbreak sizes were related to structural features of macaques' networks. For all three species, and for both anthropogenic (co-interactions) and social (grooming) contexts, outbreak sizes were positively correlated to the network centrality of first-infected macaques. Across host species and contexts, the above effects were stronger through macaques' human co-interaction networks than through their grooming networks, particularly for rhesus and bonnet macaques. Long-tailed macaques appeared to show intraspecific variation in these effects. Our findings suggest that among wildlife in anthropogenically-impacted environments, the structure of their aggregations around anthropogenic factors makes them more vulnerable to zooanthroponotic outbreaks than their social structure. The global features of these networks that influence disease outbreaks, and their underlying socio-ecological covariates, need further investigation. Animals that consistently interact with both humans and their conspecifics are important targets for disease control.

Human population expansion into nonhuman animals’ habitats has increased interest in the behavioral ecology of human-wildlife interactions. To date, however, whether and how wild animals and their conspecifics form non-random associations in terms of when or where they interact with humans still remains unclear. Here we adopt a comparative approach to address this gap, using social network analysis (SNA). SNA, increasingly implemented to determine human impact on wildlife spatial and social ecology, can be a powerful tool to understand how animal socioecology influences the spatiotemporal distribution of human-wildlife interactions. For 10 groups of rhesus, long-tailed, and bonnet macaques (Macaca spp.) living in anthropogenically-impacted environments in Asia, we collected data on human-macaque interactions, animal demographics, and macaque-macaque agonistic and affiliative social interactions. We constructed ‘human-interaction networks’ based on associations between macaques that interacted with humans within the same time and spatial locations, and social networks based on macaque-macaque allogrooming behavior, affiliative behaviors of short duration (agonistic support, lip-smacking, silent bare-teeth displays, and non-sexual mounting), and proximity. Pre-network permutation tests revealed that, for all macaque groups, human-interaction networks showed non-random structures. GLMMs revealed that individuals’ connectedness within human-interaction networks were positively associated their connectedness within affiliation social networks, and social proximity networks although this effect varied across species (bonnets > rhesus > long-tailed). Male macaques were more well-connected in human-interaction networks than females. Neither macaques’ connectedness within grooming social networks nor their dominance ranks had an impact on human-interaction networks. Our findings suggest that, in challenging, time-constraining anthropogenic environments, less time-consuming affiliative behaviors and additionally greater social tolerance (especially in less ecologically flexible species with a shorter history of exposure to human activity) may be key to animals’ maintaining strong social connections. Subsets of these animals may also utilize greater exploratory tendencies and life-histories that are less energetically demanding in the long-term. Both of these strategies may contribute to animals’ propensities to engage in joint risk-taking by being near and engaging with humans. From conservation and public health perspectives, human-interaction networks may inform interventions to mitigate zoonotic disease transmission and move human-wildlife interactions from conflict towards co-existence.

Pandemics caused by wildlife-origin pathogens, like COVID-19, highlight the importance of understanding the ecology of zoonotic transmission and outbreaks among wildlife populations at human-wildlife interfaces. To-date, the relative effects of human-wildlife and wildlife-wildlife interactions on the likelihood of such outbreaks remain unclear. In this study, we used social network analysis and epidemiological Susceptible Infected Recovered (SIR) models, to track zoonotic outbreaks through wild animals’ joint propensities to engage in social-ecological co-interactions with humans, and their social grooming interactions with conspecifics. We collected behavioral and demographic data on 10 groups of macaques ( Macaca spp.) living in (peri)urban environments across Asia. Outbreak sizes predicted by the SIR models were related to structural features of the social networks, and particular properties of individual animals’ connectivity within those networks. Outbreak sizes were larger when the first-infected animal was highly central, in both types of networks. Across host-species, particularly for rhesus and bonnet macaques, the effects of network centrality on outbreak sizes were stronger through macaques’ human co-interaction networks compared to grooming networks. Our findings, independent of pathogen-transmissibility, suggest that wildlife populations in the Anthropocene are vulnerable to zoonosis more so due to their propensities to aggregate around anthropogenic factors than their gregariousness with conspecifics. Thus, the costs of zoonotic outbreaks may outweigh the potential/perceived benefits of jointly interacting with humans to procure anthropogenic food. From One Health perspectives, animals that consistently interact with both humans and conspecifics across time and space are useful targets for disease spillover assessments and control.

Human population expansion into wildlife habitats has increased interest in the behavioural ecology of human-wildlife interactions. To date, however, the socioecological factors that determine whether, when or where wild animals take risks by interacting with humans and anthropogenic factors still remains unclear. We adopt a comparative approach to address this gap, using social network analysis (SNA). SNA, increasingly implemented to determine human impact on wildlife ecology, can be a powerful tool to understand how animal socioecology influences the spatiotemporal distribution of human-wildlife interactions. For 10 groups of rhesus, long-tailed and bonnet macaques (Macaca spp.) living in anthropogenically impacted environments in Asia, we collected data on human-macaque interactions, animal demographics, and macaque-macaque agonistic and affiliative social interactions. We constructed 'human co-interaction networks' based on associations between macaques that interacted with humans within the same time and spatial locations, and social networks based on macaque-macaque allogrooming behaviour, affiliative behaviours of short duration (agonistic support, lip-smacking, silent bare-teeth displays and non-sexual mounting) and proximity. Pre-network permutation tests revealed that, within all macaque groups, specific individuals jointly took risks by repeatedly, consistently co-interacting with humans within and across time and space. GLMMs revealed that macaques' tendencies to co-interact with humans was positively predicted by their tendencies to engage in short-duration affiliative interactions and tolerance of conspecifics, although the latter varied across species (bonnets>rhesus>long-tailed). Male macaques were more likely to co-interact with humans than females. Neither macaques' grooming relationships nor their dominance ranks predicted their tendencies to co-interact with humans. Our findings suggest that, in challenging anthropogenic environments, less (compared to more) time-consuming forms of affiliation, and additionally greater social tolerance in less ecologically flexible species with a shorter history of exposure to humans, may be key to animals' joint propensities to take risks to gain access to resources. For males, greater exploratory tendencies and less energetically demanding long-term life-history strategies (compared to females) may also influence such joint risk-taking. From conservation and public health perspectives, wildlife connectedness within such co-interaction networks may inform interventions to mitigate zoonosis, and move human-wildlife interactions from conflict towards coexistence.

Female reproductive success depends to a large extent on infants' ability to survive to maturity. While most studies of female reproductive success have focused on the effects of individuals' sociodemographic factors (e.g., age/parity, dominance rank) on offspring survival among wild primates living in less disturbed habitats, little research has focused on offspring survival in urban or periurban animals. Here we investigated sociodemographic and anthropogenic determinants of infant survival (up to 1 yr of age) in free-ranging bonnet macaques (Macaca radiata) living in a periurban environment in Southern India. We conducted the study from November 2016 to May 2018, on two groups of bonnet macaques at the Thenmala tourist site in the state of Kerala. Fifty infants were born across two birth seasons. Of these infants, 29.2% died or disappeared in 2017 and 26.9% died or disappeared in 2018. We found that infant survival was strongly influenced by the mother's parity: infants of experienced mothers had a better chance of survival than those of first-time mothers. We also found that male infants were more likely to die than female infants. However, we found no effects of mothers' dominance rank, or of frequency of mothers' interactions with humans and time spent foraging on anthropogenic food, on infant survival. Our results, consistent with findings from other wild primate species, show that even in challenging human-impacted environments, experienced bonnet macaque mothers have greater success than inexperienced ones.

Pandemics caused by wildlife-origin pathogens, like COVID-19, highlight the importance of understanding the ecology of zoonotic transmission and outbreaks among wildlife populations at human-wildlife interfaces. To-date, the relative effects of human-wildlife and wildlife-wildlife interactions on the likelihood of such outbreaks remain unclear. In this study, we used social network analysis and epidemiological Susceptible Infected Recovered (SIR) models, to track zoonotic outbreaks through wild animals joint propensities to engage in social-ecological co-interactions with humans, and their social grooming interactions with conspecifics. We collected behavioral and demographic data on 10 groups of macaques (Macaca spp.) living in (peri)urban environments across Asia. Outbreak sizes predicted by the SIR models were related to structural features of the social networks, and particular properties of individual animals connectivity within those networks. Outbreak sizes were larger when the first-infected animal was highly central, in both types of networks. Across host-species, particularly for rhesus and bonnet macaques, the effects of network centrality on outbreak sizes were stronger through macaques human co-interaction networks compared to grooming networks. Our findings, independent of pathogen-transmissibility, suggest that wildlife populations in the Anthropocene are vulnerable to zoonosis more so due to their propensities to aggregate around anthropogenic factors than their gregariousness with conspecifics. Thus, the costs of zoonotic outbreaks may outweigh the potential/perceived benefits of jointly interacting with humans to procure anthropogenic food. From One Health perspectives, animals that consistently interact with both humans and conspecifics across time and space are useful targets for disease spillover assessments and control.

OBJECTIVES In primates, allogrooming and other affiliative behaviors confer many benefits and may be influenced by many socioecological factors. Of these, the impact of anthropogenic factors remain relatively understudied. Here we ask whether interactions with humans decreased macaques' affiliative behaviors by imposing time-constraints, or increased these behaviors on account of more free-/available-time due to macaques' consumption of high-energy human foods. MATERIALS AND METHODS In Southern India, we collected data on human-macaque and macaque-macaque interactions using focal-animal sampling on two groups of semi-urban bonnet macaques for 11 months. For each macaque within each climatic season, we calculated frequencies of human-macaque interactions, rates of monitoring human activity and foraging on anthropogenic food, dominance ranks, grooming duration, number of unique grooming partners, and frequencies of other affiliative interactions. RESULTS We found strong evidence for time-constraints on grooming. Macaques that monitored humans more groomed for shorter durations and groomed fewer partners, independent of their group membership, sex, dominance rank, and season. However, monitoring humans had no impact on other affiliative interactions. We found no evidence for the free-time hypothesis: foraging on anthropogenic food was unrelated to grooming and other affiliation. DISCUSSION Our results are consistent with recent findings on other urban-dwelling species/populations. Macaques in such environments may be especially reliant on other forms of affiliation that are of short duration (e.g., coalitionary support, lip-smacking) and unaffected by time-constraints. We stress on the importance of evaluating human impact on inter-individual differences in primate/wildlife behavior for conservation efforts.

Despite increasing conflict at human–wildlife interfaces, there exists little research on how the attributes and behavior of individual wild animals may influence human–wildlife interactions. Adopting a comparative approach, we examined the impact of animals’ life-history and social attributes on interactions between humans and (peri)urban macaques in Asia. For 10 groups of rhesus, long-tailed, and bonnet macaques, we collected social behavior, spatial data, and human–interaction data for 11–20 months on pre-identified individuals. Mixed-model analysis revealed that, across all species, males and spatially peripheral individuals interacted with humans the most, and that high-ranking individuals initiated more interactions with humans than low-rankers. Among bonnet macaques, but not rhesus or long-tailed macaques, individuals who were more well-connected in their grooming network interacted more frequently with humans than less well-connected individuals. From an evolutionary perspective, our results suggest that individuals incurring lower costs related to their life-history (males) and resource-access (high rank; strong social connections within a socially tolerant macaque species), but also higher costs on account of compromising the advantages of being in the core of their group (spatial periphery), are the most likely to take risks by interacting with humans in anthropogenic environments. From a conservation perspective, evaluating individual behavior will better inform efforts to minimize conflict-related costs and zoonotic-risk.

Humans and their associated anthropogenic factors may strongly affect the demographics, activity and fragmentation of wild animal populations. Yet, the degree and nature of such impact on indicators of animals' social relationships remain largely under-investigated, despite the well-documented importance of strong social ties for an individual's health and fitness. Here, we examined whether interactions with humans may affect core aspects of social life in a primate species, by constraining the time available for individuals to engage in social interactions. Specifically, we predicted that individuals who spend more time monitoring or interacting with humans would reduce their time socializing with conspecifics (i.e. the time constraints hypothesis). Alternatively, human presence may result in increased levels of stress, leading to an increase in social behaviours that serve as a coping mechanism (i.e. the social stress hypothesis). We collected data between September 2016 and September 2017 on four groups of longtailed macaques, Macaca fascicularis, in Malaysia at two sites with differing levels of human impact. In support of the time constraints hypothesis, we found that at the site with moderate human impact, monkeys who monitored human activity more frequently engaged less often in grooming. In contrast, at the site with high human impact, we found evidence supporting the social stress hypothesis, as indicators of stress were positively associated with social interactions, although we could not link them to the presence of humans. Our results suggest that the nature of human impact on macaques' social behaviour is dependent upon the intensity of human activity and interaction with the macaques. These findings therefore provide insights into how humans may influence individual fitness and group social structure in animals living in an anthropogenic environment. More broadly, our results may lead to a better understanding of animal behaviour in anthropogenic environments, implementing conservation and population management strategies and mitigating human-wildlife conflict. (C) 2019 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

In primates, living in an anthropogenic environment can significantly improve an individual’s fitness, which is likely attributed to access to anthropogenic food resources. However, in non-professionally provisioned groups, few studies have examined whether individual attributes, such as dominance rank and sex, affect primates’ ability to access anthropogenic food. Here, we investigated whether rank and sex explain individual differences in the proportion of anthropogenic food consumed by macaques. We observed 319 individuals living in nine urban groups across three macaque species. We used proportion of anthropogenic food in the diet as a proxy of access to those food resources. Males and high-ranking individuals in both sexes had significantly higher proportions of anthropogenic food in their diets than other individuals. We speculate that unequal access to anthropogenic food resources further increases within-group competition, and may limit fitness benefits in an anthropogenic environment to certain individuals.

Time is a valuable but limited resource, and animals’ survival depends on their ability to carefully manage the amount of time they allocate to each daily activity. While existing research has examined the ecological factors affecting animals’ activity budgets, the impact of anthropogenic factors on urban-dwelling animals’ time budgets remains understudied. Here we collected data through focal animal sampling from three groups of rhesus macaques in Northern India to examine whether interactions with humans decrease macaques’ resting and social time ( time constraints hypothesis ), or whether, by contrast, foraging on anthropogenic food, that is potentially high in calories, leads macaques to spend more time resting and in social interactions ( free time hypothesis ). We found that macaques who interacted more frequently with people spent significantly less time resting and grooming, supporting the time constraints hypothesis. We argue that these time constraints are likely caused by the unpredictability of human behaviour.