Assessing dominance hierarchies: validation and advantages of progressive evaluation with Elo-rating (original) (raw)

Multicomponent and multimodal lipsmacking in crested macaques (Macaca nigra

American journal of primatology, 2013

Primates’ communicative signals are often dynamic and composed of multiple components, sometimes belonging to different sensory modalities. Such multicomponent signals are of crucial importance in the study of communication: the addition of extra features to a signal has the potential to modulate or change the meaning and message of the specific signal. Traditionally, however, components of compound signals are studied in isolation from each other, or communicative displays are studied as static and invariant wholes. Both approaches may not allow us to assess the full function of the signals. In crested macaques (Macaca nigra), the lipsmack (a display mainly used in affiliative interactions) can be produced alone or combined with other visual and acoustic communicative features. We investigated whether the composition of the lipsmack influenced the outcome of social interaction while controlling for relationship quality. Our results show that lipsmacks composed of both visual and vocal components (i.e. multimodal signals) increased the probability of affiliative contact, and therefore have enhanced signal value. The total number of visual components involved in the display had no effect but some visual components seemed more influential than others. By analyzing lipsmacking behavior as a composite, dynamic display, we were able to reveal a level of complexity that is not apparent when looking at each component separately. The results highlight the importance of a more integrative, multimodal approach to the study of primate communication. Am. J. Primatol. 75:763–773, 2013. © 2012 Wiley Periodicals, Inc.

Female parity, maternal kinship, infant age and sex influence natal attraction and infant handling in a wild colobine ( Colobus vellerosus )

American Journal of Primatology, 2014

Primate females often inspect, touch and groom others' infants (natal attraction) and they may hold and carry these infants in a manner resembling maternal care (infant handling). While natal attraction and infant handling occur in most wild colobines, little is known about the factors influencing the expression of these behaviors. We examined the effects of female parity, kinship, and dominance rank, as well as infant age and sex in wild Colobus vellerosus at Boabeng-Fiema Monkey Sanctuary, Ghana. We collected data via focal sampling of females in 2008 and 2009 (N ¼ 61) and of infants in 2010 (N ¼ 12). Accounting for the individuals who interacted with our focal subjects, this study includes 74 females and 66 infants in 8 groups. We recorded female agonistic interactions ad libitum to determine dominance ranks. We used partial pedigree information and genotypes at 17 short tandem repeat loci to determine kinship. We knew female parity, infant age and sex from demographic records. Nulliparous females showed more natal attraction and infant handling than parous females, which may suggest that interactions with infants are more adaptive for nulliparous females because they learn mothering skills through these behaviors. Compared to non-kin, maternal kin were more likely to handle infants. Maternal kin may be permitted greater access to infants because mothers are most familiar with them. Handlers may incur inclusive fitness benefits from infant handling. Dominance rank did not affect female interactions with infants. The youngest infants received the most natal attraction and infant handling, and male infants were handled more than female infants. The potential benefits of learning to mother and inclusive fitness, in combination with the relatively low costs of natal attraction and infant handling, may explain the high rates of these behaviors in many colobines. Am. J. Primatol.

GROUP DECISIONS. Shared decision-making drives collective movement in wild baboons

Science (New York, N.Y.), 2015

Conflicts of interest about where to go and what to do are a primary challenge of group living. However, it remains unclear how consensus is achieved in stable groups with stratified social relationships. Tracking wild baboons with a high-resolution global positioning system and analyzing their movements relative to one another reveals that a process of shared decision-making governs baboon movement. Rather than preferentially following dominant individuals, baboons are more likely to follow when multiple initiators agree. When conflicts arise over the direction of movement, baboons choose one direction over the other when the angle between them is large, but they compromise if it is not. These results are consistent with models of collective motion, suggesting that democratic collective action emerging from simple rules is widespread, even in complex, socially stratified societies.

Costs of and Investment in Mate-Guarding in Wild Long-Tailed Macaques (Macaca fascicularis): Influences of Female Characteristics and Male–Female Social Bonds

International Journal of Primatology, 2014

Male primates living in multimale groups tend to direct mate and mateguarding choices toward females of high reproductive value, i.e., high-ranking, parous females, or females with which they share strong bonds. Little is known, however, about the constraints that may limit male mate-guarding choices (the costs of this behavior) and the influence of the females' quality on male investment in mateguarding. We aimed to study the effects of female rank, parity status, and male-female social bond strength on the costs of and investment in mate-guarding by males. We carried out our study during two reproductive seasons on three groups of wild longtailed macaques in Indonesia. We combined behavioral observations on male locomotion and activity with noninvasive measurements of fecal glucocorticoids (fGC). Males spent less time feeding when mate-guarding nulliparous females than when mateguarding parous females and tended to have higher fGC levels when mate-guarding low-ranking nulliparous females than when mate-guarding high-ranking nulliparous ones. Evolution should thus favor male choice for high-ranking parous females because Int J Primatol such a decision brings benefits at proximate (reduced costs of mate-guarding) and ultimate (higher reproductive value) levels. Further, male investment in mate-guarding was flexible and contingent on female reproductive and social value. Males were more vigilant and more aggressive toward other males when mate-guarding females to which they were strongly bonded and/or high-ranking ones than when mate-guarding other females. Our findings bring a new dimension to the study of mate choice by showing that males not only mate preferentially with high-quality females but may also aim to secure paternity with these females through optimized monopolization.

Alpha male replacements and delayed dispersal in crested macaques ( Macaca nigra )

American Journal of Primatology, 2015

In species with a high male reproductive skew, competition between males for the top dominant position is high and escalated fights are common between competitors. As a consequence, challenges incur potentially high costs. Selection should favor males who time an alpha male challenge to maximize chances of a successful outcome minimizing costs. Despite the importance of alpha male replacements for individual males, we know little about the timing of challenges and the condition of the challenger. We investigated the timing and process of alpha male replacements in a species living in multi-male groups with high male reproductive skew, the crested macaque. We studied four wild groups over 6 years in the Tangkoko Reserve, North Sulawesi, Indonesia, during which 16 alpha male replacements occurred. Although unusual for cercopithecines, male crested macaques delayed their natal dispersal until they attained maximum body mass and therefore fighting ability whereupon they emigrated and challenged the alpha male in another group. Accordingly, all observed alpha male replacements were from outside males. Ours is the first report of such a pattern in a primate species living in multi-male groups. Although the majority of alpha male replacements occurred through direct male-male challenges, many also took place opportunistically (i.e., after the alpha male had already been injured or had left the group). Furthermore, alpha male tenures were very short (averaging ca. 12 months). We hypothesize that this unusual pattern of alpha male replacements in crested macaques is related to the species-specific combination of high male reproductive skew with a large number of males per group. Am. J. Primatol.

A comparison of dominance rank metrics reveals multiple competitive landscapes in an animal society

Proceedings of the Royal Society B: Biological Sciences

Across group-living animals, linear dominance hierarchies lead to disparities in access to resources, health outcomes and reproductive performance. Studies of how dominance rank predicts these traits typically employ one of several dominance rank metrics without examining the assumptions each metric makes about its underlying competitive processes. Here, we compare the ability of two dominance rank metrics—simple ordinal rank and proportional or ‘standardized’ rank—to predict 20 traits in a wild baboon population in Amboseli, Kenya. We propose that simple ordinal rank best predicts traits when competition is density-dependent, whereas proportional rank best predicts traits when competition is density-independent. We found that for 75% of traits (15/20), one rank metric performed better than the other. Strikingly, all male traits were best predicted by simple ordinal rank, whereas female traits were evenly split between proportional and simple ordinal rank. Hence, male and female tra...

Multimodal Advertisement of Pregnancy in Free-Ranging Female Japanese Macaques (Macaca fuscata)

PLOS ONE, 2015

The role of multiple sexual signals in indicating the timing of female ovulation, and discrimination of this timing by males, has been particularly well studied among primates. However the exhibition of pregnancy signals, and how such signals might modulate male post-conception mating decisions, is still poorly understood. Here we aimed to determine if Japanese macaque males use changes in female sexual signals (behavioral, visual and auditory) to discriminate pregnancy and adjust their socio-sexual behaviors. We combined behavioral observations, digital photography and endocrinological (progestogen and estrogen) data, collected systematically during three one-month periods: the pre-conceptive period, the 1 st month of pregnancy and the 2 nd month of pregnancy. We analyzed variation in the probability of detecting male and female socio-sexual behaviors and estrus calls, as well as changes in female face color parameters, in relation to female reproductive state. Based on our focal observations, we found that males did not copulate during the pregnancy period, and that female socio-sexual behaviors generally decreased from the pre-conceptive to post-conceptive periods. Female face luminance decreased from the pre-conceptive month to the pregnancy period whereas face color only varied between the 1 st and 2 nd month of gestation. Our results suggest that Japanese macaque females display sexual cues of pregnancy that males might use to reduce energy wasted on non-reproductive copulations with pregnant females. We hypothesize that females advertize their pregnancy through changes in behavioral, visual and potential auditory signals that males can use to adjust their mating behaviors. We finish by discussing implications for male and female post-conception strategies.

Reestablishment of Social Hierarchies in Weaned Pigs after Mixing

Animals

Pigs are animals that live in groups and have social hierarchies within the group. After mixing, they can re-establish social hierarchies within several days through fighting. Dominance hierarchical indices, such as I&SI, Elo rating, and Glicko rating, have been used to analyze social hierarchies of some social animals but not pigs. I&SI index involves iterative calculations that first minimize the number of inconsistencies (I) in a dominance matrix, and then minimize the strength of those inconsistencies (SI). Elo rating and Glicko rating indices are based on the sequence in which interactions occur, and continuously update ratings by looking at interactions sequentially. To study the temporal dynamics of social hierarchy formation and maintenance in weaned pigs after mixing, a total of 102 pigs (47 females and 55 barrows) were selected by similar body weight and mixed in 10 pens (10 or 11 females or barrows per pen). Their behavior was recorded and observed for 72 h after mixing. ...

Social thermoregulation as a potential mechanism linking sociality and fitness: Barbary macaques with more social partners form larger huddles

Scientific reports, 2018

Individuals with more or stronger social bonds experience enhanced survival and reproduction in various species, though the mechanisms mediating these effects are unclear. Social thermoregulation is a common behaviour across many species which reduces cold stress exposure, body heat loss, and homeostatic energy costs, allowing greater energetic investment in growth, reproduction, and survival, with larger aggregations providing greater benefits. If more social individuals form larger thermoregulation aggregations due to having more potential partners, this would provide a direct link between sociality and fitness. We conducted the first test of this hypothesis by studying social relationships and winter sleeping huddles in wild Barbary macaques (Macaca sylvanus), wherein individuals with more social partners experience greater probability of winter survival. Precipitation and low temperature increased huddle sizes, supporting previous research that huddle size influences thermoregul...

Matrilineal Behavioral and Physiological Changes following the Death of a Non-Alpha Matriarch in Rhesus Macaques (Macaca mulatta)

PLOS ONE, 2016

In many species, the loss of alpha matriarchs is associated with a number of negative outcomes such as troop fission, eviction, wounding, and reduced vitality. However, whether the dramatic consequences of their loss are due to their role as an old experienced figure or to their alpha status remains unclear. In a retrospective study, we tested that in a semi-free ranging colony of rhesus macaques (Macaca mulatta), the removal of a non-alpha matriarch, who had a large set of kin, led to changes in behavior and physiological stress within her matriline. Following her removal, her matriline increased in aggression, vigilance, and social grooming. Additionally, hierarchical stability, measured by levels of rank changes, decreased within her matriline, and levels of intense aggression by high-ranking animals were more frequent, as well as matrilineal wounding. Although ordinal rank was positively associated with higher chronic hair cortisol concentrations (HCCs) in the months before the matriarch's removal, following her removal, only those who experienced large increases in rank within her matriline displayed higher HCCs. Changes in matrilineal stability, aggression, behavior, and HCCs within the other two matrilines in the troop were not evident, although caution is needed due to the small sample sizes. We conclude that the removal of the non-alpha matriarch led to matrilineal instability, characterized by higher levels of aggression and subsequent vigilance, rank changes, physiological stress, and grooming. We suggest that non-alpha matriarchs with a large number of kin and social support can be integral to the stability of matrilines.

Social groups buffer maternal loss in mountain gorillas

eLife

Mothers are crucial for mammals’ survival before nutritional independence, but many social mammals reside with their mothers long after. In these species the social adversity caused by maternal loss later in life can dramatically reduce fitness. However, in some human populations these negative consequences can be overcome by care from other group members. We investigated the consequences of maternal loss in mountain gorillas and found no discernible fitness costs to maternal loss through survival, age at first birth, or survival of first offspring through infancy. Social network analysis revealed that relationships with other group members, particularly dominant males and those close in age, strengthened following maternal loss. In contrast to most social mammals, where maternal loss causes considerable social adversity, in mountain gorillas, as in certain human populations, this may be buffered by relationships within cohesive social groups, breaking the link between maternal loss...

Neophobia in 10 ungulate species—a comparative approach

Behavioral Ecology and Sociobiology

Neophobia (the fearful reaction to novel stimuli or situations) has a crucial effect on individual fitness and can vary within and across species. However, the factors predicting this variation are still unclear. In this study, we assessed whether individual characteristics (rank, social integration, sex) and species socio-ecological characteristics (dietary breadth, group size, domestication) predicted variation in neophobia. For this purpose, we conducted behavioral observations and experimental tests on 78 captive individuals belonging to 10 different ungulate species—an ideal taxon to study inter-specific variation in neophobia given their variety in socio-ecological characteristics. Individuals were tested in their social groups by providing them with familiar food, half of which had been positioned close to a novel object. We monitored the individual latency to approach and eat food and the proportion of time spent in its proximity. Using a phylogenetic approach and social net...

Dominance style only partially predicts differences in neophobia and social tolerance over food in four macaque species

Scientific Reports

Primates live in complex social systems with social structures ranging from more to less despotic. In less despotic species, dominance might impose fewer constraints on social choices, tolerance is greater than in despotic species and subordinates may have little need to include novel food items in the diet (i.e. neophilia), as contest food competition is lower and resources more equally distributed across group members. Here, we used macaques as a model to assess whether different dominance styles predict differences in neophilia and social tolerance over food. We provided familiar and novel food to 4 groups of wild macaques (N = 131) with different dominance styles (Macaca fuscata, M. fascicularis, M. sylvanus, M. maura). Our study revealed inter- and intra-specific differences in individuals’ access to food, which only partially reflected the dominance styles of the study subjects. Contrary to our prediction, social tolerance over food was higher in more despotic species than in ...

Comparative Investigations of Social Context-Dependent Dominance in Captive Chimpanzees (Pan troglodytes) and Wild Tibetan Macaques (Macaca thibetana)

Scientific reports, 2018

Theoretical definitions of dominance, how dominance is structured and organized in nature, and how dominance is measured have varied as investigators seek to classify and organize social systems in gregarious species. Given the variability in behavioral measures and statistical methods used to derive dominance rankings, we conducted a comparative analysis of dominance using existing statistical techniques to analyze dominance ranks, social context-dependent dominance structures, the reliability of statistical analyses, and rank predictability of dominance structures on other social behaviors. We investigated these topics using behavioral data from captive chimpanzees (Pan troglodytes) and wild Tibetan macaques (Macaca thibetana). We used a combination of all-occurrence, focal-animal, and instantaneous scan sampling to collect social, agonistic, and associative data from both species. We analyzed our data to derive dominance ranks, test rank reliability, and assess cross-context pred...

Oxytocin reactivity during intergroup conflict in wild chimpanzees

Proceedings of the National Academy of Sciences, 2016

Intergroup conflict is evident throughout the history of our species, ubiquitous across human societies, and considered crucial for the evolution of humans’ large-scale cooperative nature. Like humans, chimpanzee societies exhibit intragroup coordination and coalitionary support during violent intergroup conflicts. In both species, cooperation among group members is essential for individuals to gain access to benefits from engaging in intergroup conflict. Studies suggest that a contributive mechanism regulating in-group cooperation during intergroup conflicts in humans involves the neuropeptide hormone oxytocin, known to influence trust, coordination, and social cognition, although evidence from natural settings is lacking. Here, applying a noninvasive method, we investigate oxytocinergic system involvement during natural intergroup conflicts in wild chimpanzees. We found that chimpanzees of both sexes had significantly higher urinary oxytocin levels immediately before and during in...

Bystanders intervene to impede grooming in Western chimpanzees and sooty mangabeys

Royal Society open science, 2017

Grooming interactions benefit groomers, but may have negative consequences for bystanders. Grooming limits bystanders' grooming access and ensuing alliances could threaten the bystander's hierarchy rank or their previous investment in the groomers. To gain a competitive advantage, bystanders could intervene into a grooming bout to increase their own grooming access or to prevent the negative impact of others' grooming. We tested the impact of dominance rank and social relationships on grooming intervention likelihood and outcome in two sympatric primate species, Western chimpanzees () and sooty mangabeys (). In both species, rather than increasing their own access to preferred partners, bystanders intervened mainly when an alliance between groomers could have a negative impact on them: when the lower-ranking groomer was close to the bystander in rank, when either groomer was an affiliation partner whose services they could lose, or the groomers were not yet strongly affi...

Beyond the Energy Poor/Non Energy Poor Divide: Energy Vulnerability and Mindsets on Energy Generation Modes in Hungary

Energies

In the Hungarian as well as the international literature, debates regarding the methodology used for measuring energy poverty are ongoing. Our contribution sought to develop a methodology that would be relatively easy to measure via survey and have the potential to identify households’ energy vulnerability status beyond the energy poor/non energy poor divide. Based on a representative survey that was carried out in Hungary in 2018, we identified three groups of households: energy poor, non energy poor households, and a so-called transitional group, potentially at risk of energy poverty. In addition to socioeconomic differences, we analysed differences in the mindsets of respondents belonging to these three household groups concerning different aspects of heating energy generation (macrolevel aspects, such as environmental sustainability and energy supply system issues, and microlevel aspects, such as economic and technical issues). By this, the article aims to provide input for furt...

Dominance style predicts differences in food retrieval strategies

Scientific Reports

In several species, rank predicts access to food, and subordinates may need specific behavioural strategies to get a share of resources. This may be especially important in despotic species, where resources are strongly biased in favour of dominants and subordinates may more strongly rely on specific tactics to maximize food intake. Here, we compared three macaque species with an experimental set-up reproducing feeding competition contest. Following our predictions, more tolerant species mostly retrieved food in the presence of others and were less dependent on specific tactics. Contrarily, subordinates in more despotic species more likely collected food (1) when dominants could not see food or (2) were attacking others, (3) while “dissimulating”, or (4) “storing food”. Our study reveals that dominance styles reliably predict the probability of using specific food retrieval tactics and provides important insights on the social conditions that might have led to the emergence of tacti...

Group housing and social dominance hierarchy affect circadian activity patterns in mice

Royal Society Open Science

In this study, we investigated the effect of social environment on circadian patterns in activity by group housing either six male or six female mice together in a cage, under regular light–dark cycles. Based on the interactions among the animals, the social dominance rank of individual mice was quantitatively established by calculating Elo ratings. Our results indicated that, during our experiment, the social dominance hierarchy was rapidly established, stable yet complex, often showing more than one dominant mouse and several subordinate mice. Moreover, we found that especially dominant male mice, but not female mice, displayed a significantly higher fraction of their activity during daytime. This resulted in reduced rhythm amplitude in dominant males. After division into separate cages, male mice showed an enhancement of their 24 h rhythm, due to lower daytime activity. Recordings of several physiological parameters showed no evidence for reduced health as a potential consequence...

Personality of Wild Male Crested Macaques (Macaca nigra)

PLoS ONE, 2013

Animal personalities, i.e. consistent differences in behavior across time and/or context, have received increased attention of behavioral biologists over the last years. Recent research shows that personalities represent traits on which natural and sexual selection work and which can have substantial fitness consequences. The aim of this study is to establish the personality structure of crested macaque (Macaca nigra) males as foundation for future studies on its adaptive value. We collected behavioral data through focal animal sampling and additionally conducted two sets of playback experiments. Results of a factor analysis on the behavioral data revealed a four factor structure with components we labeled Anxiety, Sociability, Connectedness and Aggressiveness. Results from the experiments revealed an additional and independent Boldness factor but the absence of Neophilia. Overall, this structure resembles other macaque and animal species with the exception of Connectedness, which might be a consequence of the species' tolerant social style. Our results thus not only form the basis for future studies on the adaptive value of personality in crested macaques but also contribute an important data point for investigating the evolution of personality structure from a comparative perspective by refining, for example, which personality factors characterized the last common ancestor of hominids and macaques.

Female fertile phase synchrony, and male mating and reproductive skew, in the crested macaque

Scientific Reports

High social status is the primary determinant of reproductive success among group-living male mammals. Primates living in multimale–multifemale groups show the greatest variation in the strength of this link, with marked variation in reproductive skew by male dominance among species, dependent on the degree of female fertile phase synchrony, and the number of competing males. Here, we present data on two groups of wild crested macaques (Macaca nigra), living in the Tangkoko Reserve, Sulawesi, Indonesia. We investigated male monopolization of fertile females in 31 cycles of 19 females, and genetic paternity of 14 offspring conceived during the study period. We show that female fertile phase synchrony was low, that females had few mating partners in their fertile phase, and that dominant males monopolized a high proportion of consortships and matings, resulting in marked and steep mating and reproductive skew. We conclude that female cycle asynchrony provides the opportunity for stron...

Oil palm cultivation critically affects sociality in a threatened Malaysian primate

2021

Human-induced habitat alterations globally threaten animal populations, often evoking complex behavioural responses in wildlife. This may be particularly dramatic when negatively affecting social behaviour, which fundamentally determines individual fitness and offspring survival in group-living animals. Here, we provide first evidence for significant behavioural modifications in sociality of southern pig-tailed macaques visiting Malaysian oil palm plantations in search of food despite elevated predation risk. Specifically, we found critical reductions of key positive social interactions but higher rates of aggression in the plantation interior compared to the plantation edge (i.e. plantation areas bordering the forest) and the forest. At the plantation edge, affiliation even increased compared to the forest, while central positions in the macaques' social network structure shifted from high-ranking adult females and immatures to low-ranking individuals. Further, plantations also...

Female preferences for male golden snub-nosed monkeys vary with male age and social context

Current Zoology

Age is a key factor affecting sexual selection, as many physical and social traits are age-related. Although studies of primate mate choice often consider particular age-related traits, few consider the collective effects of male age. We tested the hypothesis that female golden snub-nosed monkeys Rhinopithecus roxellana prefer prime aged males (10–15 years) over younger and older males. We examined a habituated, provisioned troop during a 3-year study in the Qinling Mountains, China. Prime age males were more likely to be resident males of 1-male units (OMUs) than males of other ages. Since females are free to transfer between OMUs, the number of females per OMU can be indicative of female preferences. We examined the number of females per OMU, and found that it increased with resident male age up to 7–8 years, and declined after 12 years, such that prime age resident males had more females than other resident males. Females also initiated extra-unit copulations with high-ranking pr...

The effect of individual and food characteristics on food retrieval and food sharing in captive Guinea baboons ( Papio papio )

American Journal of Primatology

Access to food is of major importance to the fitness and survival of every individual, particularly in group-living animals, in which individual characteristics and food distribution can affect food intake. Additionally, several species of primates are known to share food under certain conditions. Such unresisted transfer of food from one individual to another appears to be adaptive, for instance as a tool to maintain and reinforce social bonds. In this study, we aimed to test how food retrieval and food sharing varies depending on the social relationship between individuals, and on the characteristics of the food. In six different test conditions, we provided a captive group of Guinea baboons (Papio papio, N = 23) with multiple food items, differing in quality, quantity, density, monopolizability, and effort required to obtain it. We further used behavioral observations to assess individual relationships and possible variations in grooming exchanges linked to food sharing events. Out of 424 events in which food items were retrieved by the subjects, we detected no instances of active food sharing and only 17 of passive food sharing. The way food was retrieved was affected by individual and food characteristics (i.e., quantity, quality, and monopolizability of food): Males and central individuals (i.e., those connected to many partners, and/or having partners with many connections in the social network) were more likely to retrieve food during test conditions. In particular, events of passive food sharing mostly happened when the quality of food was low, and between individuals belonging to the same community (i.e., having close relationships). No other food characteristics affected the probability to share food, and the occurrence of food sharing had no immediate effect on grooming exchanges. Overall, our findings suggest that food sharing is relatively rare in Guinea baboons unless the food has a low quality and individuals form close social bonds.

Behavioural patterns of vocal greeting production in four primate species

Royal Society Open Science

Social animals have evolved a range of signals to avoid aggressive and facilitate affiliative interactions. Vocal behaviour is especially important in this respect with many species, including various primates, producing acoustically distinct ‘greeting calls’ when two individuals approach each other. While the ultimate function of greeting calls has been explored in several species, little effort has been made to understand the mechanisms of this behaviour across species. The aim of this study was to explore how differences in individual features (individual dominance rank), dyadic relationships (dominance distance and social bond strength) and audience composition (presence of high-ranking or strongly bonded individuals in proximity), related to vocal greeting production during approaches between two individuals in the philopatric sex of four primate species: female olive baboons ( Papio anubis ), male chimpanzees ( Pan troglodytes ), female sooty mangabeys ( Cercocebus atys ) and ...

Sequential information in a great ape utterance

Scientific Reports

Birdsong is a prime example of acoustically sophisticated vocal behaviour, but its complexity has evolved mainly through sexual selection to attract mates and repel sexual rivals. In contrast, non-human primate calls often mediate complex social interactions, but are generally regarded as acoustically simple. Here, we examine arguably the most complex call in great ape vocal communication, the chimpanzee (Pan troglodytes schweinfurthii) 'pant hoot'. This signal consists of four acoustically distinct phases: introduction, build-up, climax and let-down. We applied state-of-theart Support Vector Machines (SVM) methodology to pant hoots produced by wild male chimpanzees of Budongo Forest, Uganda. We found that caller identity was apparent in all four phases, but most strongly in the low-amplitude introduction and high-amplitude climax phases. Age was mainly correlated with the low-amplitude introduction and build-up phases, dominance rank (i.e. social status) with the high-amplitude climax phase, and context (reflecting activity of the caller) with the lowamplitude let-down phase. We conclude that the complex acoustic structure of chimpanzee pant hoots is linked to a range of socially relevant information in the different phases of the call, reflecting the complex nature of chimpanzee social lives.

Data-driven team ranking and match performance analysis in Chinese Football Super League

Chaos, Solitons & Fractals

Recent years have seen an increasing body of research into the evaluation of the team-level technicaltactical performance in association football using match events data. However, most studies used monodimensional approach and modeled the influence of each performance aspects on match result in isolation, which limited the interpretability of the results. The study was aimed to apply a state-of-the-art algorithm to the ranking of team performance and exploitation of key performance features in relation to match outcome based on massive match dataset. Data of all 1200 matches from 2014 to 2018 Chinese Football Super League (CSL) were used. From the original 164 match events, we extracted 22 features that were related to attacking, passing, and defending performance and most. A Linear Support Vector Classifier (LSVC) model was subsequently built with these 22 input features and trained in order to rank the teams by their performance and analyze the features that influence most match outcome (win/not win), with the dataset being divided into a ratio of 4:1 to train and validate the model. The results have shown that the data-driven LSVC model displayed a prediction accuracy of 0.83 and the ranking of teams' match performance and prediction of teams' league standings were highly correlated with their actual ranking. Saves, pass success and shot on target in penalty area were demonstrated as top positive features for winning whereas shots on target during open play, pass and bad shot% were three negative features most influential for the match result. The team ranks of all teams were highly correlated with their real final league rankings. In general, CSL winning teams build their success based on defensive ability and shooting accuracy, and high-ranked teams could always maintain better performance than their counterparts. The team-rank framework could provide a consolidated and complex approach to evaluate the match performance quality of the teams, refining decisions-making during match preparation and player transfer at different periods of the season.

Social status alters immune regulation and response to infection in macaques

Science, 2016

Status alters immune function in macaques Rhesus macaques experience variable levels of stress on the basis of their position in the social hierarchy. To examine how stress affects immune function, Snyder-Mackler et al. manipulated the social status of individual macaques (see the Perspective by Sapolsky). Social status influenced the immune system at multiple levels, from immune cell numbers to gene expression, and altered signaling pathways in a model of response to infection. Macaques possess a plastic and adaptive immune response wherein social subordination promotes antibacterial responses, whereas high social status promotes antiviral responses. Science , this issue p. 1041 ; see also p. 967

Social status alters chromatin accessibility and the gene regulatory response to glucocorticoid stimulation in rhesus macaques

Proceedings of the National Academy of Sciences

Low social status is an important predictor of disease susceptibility and mortality risk in humans and other social mammals. These effects are thought to stem in part from dysregulation of the glucocorticoid (GC)-mediated stress response. However, the molecular mechanisms that connect low social status and GC dysregulation to downstream health outcomes remain elusive. Here, we used an in vitro GC challenge to investigate the consequences of experimentally manipulated social status (i.e., dominance rank) for immune cell gene regulation in female rhesus macaques, using paired control and GC-treated peripheral blood mononuclear cell samples. We show that social status not only influences immune cell gene expression but also chromatin accessibility at hundreds of regions in the genome. Social status effects on gene expression were less pronounced following GC treatment than under control conditions. In contrast, social status effects on chromatin accessibility were stable across conditi...

Thermal imaging reveals audience-dependent effects during cooperation and competition in wild chimpanzees

Scientific Reports

Accessing animal minds has remained a challenge since the beginnings of modern science. Here, we used a little-tried method, functional infrared thermal imaging, with wild chimpanzees during common social interactions. After removing confounds, we found that chimpanzees involved in competitive events had lower nose skin temperatures whereas those involved in cooperative events had higher temperatures, the latter more so in high- than low-ranking males. Temperatures associated with grooming were akin to those of cooperative events, except when males interacted with a non-reciprocating alpha male. In addition, we found multiple audience effects. Notably, the alpha male’s presence reduced positive effects associated with cooperation, whereas female presence buffered negative effects associated with competition. Copulation was perceived as competitive, especially during furtive mating when other males were absent. Overall, patterns suggest that chimpanzees categorise ordinary social eve...

Maternal effects on early-life gut microbiome maturation in a wild nonhuman primate

2021

ABSTRACTEarly-life gut microbial colonization is an important process shaping host physiology, immunity and long-term health outcomes in humans and other animals. However, our understanding of this dynamic process remains poorly investigated in wild animals, where developmental mechanisms can be better understood within ecological and evolutionary relevant contexts. Using 16s rRNA amplicon sequencing on 525 fecal samples from a large cohort of infant and juvenile geladas (Theropithecus gelada), we characterized gut microbiome maturation during the first three years of life and assessed the role of maternal effects in shaping offspring microbiome assembly. Microbial diversity increased rapidly in the first months of life, followed by more gradual changes until weaning. As expected, changes in gut microbiome composition and function with increasing age reflected progressive dietary transitions: in early infancy when infants rely heavily on their mother’s milk, microbes that facilitate...

Concede or clash? Solitary sharks competing for food assess rivals to decide

Proceedings of the Royal Society B: Biological Sciences, 2018

To adapt to their environment, organisms can either directly interact with their surroundings or use social information (i.e. information provided by neighbouring individuals). Social information relates to the external features of surrounding peers, and little is known about its use by solitary species. Here, we investigated the use of social cues in a solitary marine predator by creating artificial aggregations of free-ranging sicklefin lemon sharks ( Negaprion acutidens ). Using a novel monitoring protocol, we analysed both dominance interactions and tolerance associations between sharks competing for food in relation with the number, the morphology and the behaviour of rivals. Sharks produced more agonistic displays and spent more time around the bait as competitors were more abundant. Moreover, the morphological attributes of competitors had very limited influence on the structure of shark social interactions. Instead, sharks appeared to establish tolerance relationships with c...

Energetic management in wild chimpanzees (Pan troglodytes verus) in Taï National Park, Côte d’Ivoire

Behavioral Ecology and Sociobiology, 2020

Socioecological theories predict that, in mammals, feeding and mating competitions affect male and female energetic conditions differently but energetic studies investigating both sexes simultaneously are rare. We investigated the effect of socioecological factors on the energetic conditions of male and female western chimpanzees, a long-lived species with high degrees of male-male competition. We used behavioural data collected on one chimpanzee community in the Taï National Park over 12 months, phenological data and urinary c-peptide (UCP) measures, a marker of energy balance. We found a positive effect of food availability on UCP levels in both sexes. Dominance rank also affected chimpanzee UCP levels. High-ranking females had higher UCP levels than low-ranking ones but only in periods when no oestrus females were present in the community. In contrast, high-ranking males had higher UCP levels than low-ranking males in the presence of oestrus females but lower UCP levels in their ...

Evidence of joint commitment in great apes' natural joint actions

Royal Society Open Science, 2021

Human joint action seems special, as it is grounded in joint commitment—a sense of mutual obligation participants feel towards each other. Comparative research with humans and non-human great apes has typically investigated joint commitment by experimentally interrupting joint actions to study subjects’ resumption strategies. However, such experimental interruptions are human-induced, and thus the question remains of how great apes naturally handle interruptions. Here, we focus on naturally occurring interruptions of joint actions, grooming and play, in bonobos and chimpanzees. Similar to humans, both species frequently resumed interrupted joint actions (and the previous behaviours, like grooming the same body part region or playing the same play type) with their previous partners and at the previous location. Yet, the probability of resumption attempts was unaffected by social bonds or rank. Our data suggest that great apes experience something akin to joint commitment, for which w...

Rapid transmission of respiratory infections within but not between mountain gorilla groups

Scientific Reports, 2021

Minimizing disease transmission between humans and wild apes and controlling outbreaks in ape populations is vital to both ape conservation and human health, but information on the transmission of real infections in wild populations is rare. We analyzed respiratory outbreaks in a subpopulation of wild mountain gorillas (Gorilla beringei beringei) between 2004 and 2020. We investigated transmission within groups during 7 outbreaks using social networks based on contact and proximity, and transmission between groups during 15 outbreaks using inter-group encounters, transfers and home range overlap. Patterns of contact and proximity within groups were highly predictable based on gorillas’ age and sex. Disease transmission within groups was rapid with a median estimated basic reproductive number (R0) of 4.18 (min = 1.74, max = 9.42), and transmission was not predicted by the social network. Between groups, encounters and transfers did not appear to have enabled disease transmission and ...

Correlates of androgens in wild male Barbary macaques: Testing the challenge hypothesis

American journal of primatology, 2017

Investigating causes and consequences of variation in hormonal expression is a key focus in behavioral ecology. Many studies have explored patterns of secretion of the androgen testosterone in male vertebrates, using the challenge hypothesis (Wingfield, Hegner, Dufty, & Ball, 1990; The American Naturalist, 136(6), 829-846) as a theoretical framework. Rather than the classic association of testosterone with male sexual behavior, this hypothesis predicts that high levels of testosterone are associated with male-male reproductive competition but also inhibit paternal care. The hypothesis was originally developed for birds, and subsequently tested in other vertebrate taxa, including primates. Such studies have explored the link between testosterone and reproductive aggression as well as other measures of mating competition, or between testosterone and aspects of male behavior related to the presence of infants. Very few studies have simultaneously investigated the links between testoste...

Repeatable glucocorticoid expression is associated with behavioural syndromes in males but not females in a wild primate

Royal Society Open Science, 2019

Behavioural syndromes are a well-established phenomenon in human and non-human animal behavioural ecology. However, the mechanisms that lead to correlations among behaviours and individual consistency in their expression at the apparent expense of behavioural plasticity remain unclear. The ‘state-dependent' hypothesis posits that inter-individual variation in behaviour arises from inter-individual variation in state and that the relative stability of these states within an individual leads to consistency of behaviour. The endocrine stress response, in part mediated by glucocorticoids (GCs), is a proposed behavioural syndrome-associated state as GC levels are linked to an individual's behavioural responses to stressors. In this study, in wild Barbary macaques ( Macaca sylvanus ), consistent inter-individual differences were observed in both sexes for GC activity (faecal glucocorticoid, fGC concentrations), but not GC variation (coefficient of variation in fGC concentrations)....

Chimpanzee lip-smacking facilitates cooperative behaviour

Scientific reports, 2015

Signalling plays an important role in facilitating and maintaining affiliative or cooperative interactions in social animals. Social grooming in primates is an example of an interaction that requires coordination between partners but little is known about communicative behaviours facilitating this activity. In this study, we analysed the communication of wild chimpanzees of Budongo Forest, Uganda, as they entered and maintained a naturally occurring cooperative interaction: social grooming. We found that lip-smacking, a distinct multimodal oral gesture produced during grooming, coordinated this activity. Lip-smacking at the beginning of grooming bouts was significantly more often followed by longer and reciprocated bouts than silent grooming initiations. Lip-smacks were more likely to be produced when the risk of termination of the interaction by the recipient was high, for instance when grooming vulnerable body parts. Groomers were also more likely to produce lip-smacks during face...

An Audience Effect in Sooty Mangabey Alarm Calling

Frontiers in Psychology, 2022

How does intentional communication evolve? Comparative studies can shed light on the evolutionary history of this relevant feature of human language and its distribution before modern humans. The current animal literature on intentional signaling consists mostly of ape gestural studies with evidence of subjects persisting and elaborating with sometimes arbitrary signals toward a desired outcome. Although vocalizations can also have such imperative qualities, they are typically produced in a functionally fixed manner, as if evolved for a specific purpose. Yet, intentionality can sometimes transpire even in functionally fixed calls, for example, if production is adapted to audience composition. In this study, we carried out field experiments to test whether free-ranging sooty mangabeys adjusted snake alarm call production to their audiences. We found a positive relation between alarm call production and naïve individuals arriving, suggesting that callers attempted to influence their b...

Habitat Quality and Social Behavioral Association Network in a Wintering Waterbirds Community

Sustainability

Migratory waterbirds concentrated in freshwater ecosystems in mosaic environments rely on quality habitats for overwintering. At West Dongting Lake National Nature Reserve (WDLNNR), China, land-use change and hydrology alternation are compounding factors that have affected important wintering areas for migratory waterbirds. Presently, changes in the hydrology and landscape have reshaped natural wintering habitats and their availability, though the impact of hydrological management on habitat selection of wintering waterbirds is largely unknown. In this study, we classified differentially managed habitats and calculated their area using the normalized difference vegetation index (NDVI) to evaluate suitable habitat availability over the study period (2016–2017 and 2017–2018 wintering periods). We then used social behavioral association network (SBAN) model to compare habitat quality through species-species social interactions and species-habitat associations in lakes with different hy...

Construct social‐behavioral association network to study management impact on waterbirds community ecology using digital video recording cameras

Ecology and Evolution, 2021

Studying social‐behavior and species associations in ecological communities is challenging because it is difficult to observe the interactions in the field. Animal behavior is especially difficult to observe when selection of habitat and activities are linked to energy costs of long‐distance movement. Migrating communities tend to be resource specific and prefer environments that offer more suitability for coexisting in a shared space and time. Given the recent advances in digital technologies, digital video recording systems are gaining popularity in wildlife research and management. We used digital video recording cameras to study social interactions and species–habitat linkages for wintering waterbirds communities in shared habitats. Examining over 8,640 hr of video footages, we built tetrapartite social‐behavioral association network of wintering waterbirds over habitat (n = 5) selection events in sites with distinct management regimes. We analyzed these networks to identify hub...

Informed horses are influential in group movements, but they may avoid leading

Animal Cognition, 2015

In species that move in cohesive groups, animals generally reach decisions through socially distributed processes, and individual knowledge is expected to influence collective decisionmaking. The pooling of information should not be considered a general rule, however, since conflicts of interest may occur between group members. When resources are limited or highly attractive, higher-ranking individuals can prevent others from accessing food, and subordinates may have an interest in withholding information its location. We investigated the role individual knowledge may play in recruitment processes in four groups of horses (Equus caballus). Animals were repeatedly released in a food search situation, in which one individual had been informed about the location of a preferred food, while another was a naïve control subject. Horses that were informed about the location of the feeding site were seen to approach the food source more steadily, and were followed by a higher number of group members than their uninformed counterparts. Recruitment processes appeared mostly passive. Amongst the informed subjects, lower-ranking individuals were overall less followed than the higher-ranking ones. Most lower-ranking horses arrived alone at the feeding site. Non-followed informed subjects spent less time in active walk and used direct paths less frequently than followed animals, they were joined by fewer partners at the attractive food source and spent more time feeding alone. This indicates that the influence of informed individuals on the behavior of other group members was a mixed process. Some horses brought nutritional benefits to their conspecifics by leading them to food supplies, whereas the behavior of others might be functionally deceptive.

Think Fast!: Vervet Monkeys Assess the Risk of Being Displaced by a Dominant Competitor When Making Foraging Decisions

Frontiers in Ecology and Evolution

Foraging animals need to quickly assess the costs and benefits of different foraging decisions, including resource quantity, quality, preference, ease of access, dispersion, distance, and predation risk. Social animals also need to take social context into account and adapt foraging strategies that maximize net resource intake and minimize contest competition with conspecifics. We used an experimental approach to investigate how social context impacts wild vervet monkey (Chlorocebus pygerythrus) foraging decisions in a multi-destination pentagon array. We baited four platforms with less-preferred corn and one platform with a larger, preferred resource (half banana) that required handling time. We ran over 1,000 trials and found that when monkeys foraged alone, they usually took the path that minimized travel distance but prioritized the preferred-food platform when in competition. However, the foraging strategy chosen by low-ranking individuals depended on the handling skill of the ...

Social style and resilience of macaques’ networks, a theoretical investigation

Primates, 2018

Group-living animals rely on efficient transmission of information for optimal exploitation of their habitat. How efficient and resilient a network is depend on its structure, which is a consequence of the social interactions of the individuals that comprise the network. In macaques, network structure differs according to dominance style. Networks of intolerant species are more modular, more centralized, and less connected than those of tolerant ones. Given these structural differences, networks of intolerant species are potentially more vulnerable to fragmentation and decreased information transmission when central individuals disappear. Here we studied network resilience and efficiency in artificial societies of macaques. The networks were produced with an individual-based model that has been shown to reproduce the structural features of networks of tolerant and intolerant macaques. To study network resilience, we deleted either central individuals or individuals at random and studied the effects of these deletions on network cohesiveness and efficiency. The deletion of central individuals had more negative effects than random deletions from the networks of both tolerant and intolerant artificial societies. Central individuals thus appeared to aid in the maintenance of network cohesiveness and efficiency. Further, the networks of both intolerant and tolerant societies appeared to be robust to the loss of individuals, as network fragmentation was never observed. Our results suggest that despite differences in network structure, networks of tolerant and intolerant macaques may be equally resilient.

Long term analysis of social structure: evidence of age-based consistent associations in male Alpine ibex

2021

ABSTRACTDespite its recognized importance for understanding the evolution of animal sociality as well as for conservation, long term analysis of social networks of animal populations is still relatively uncommon. We investigated social network dynamics in males of a gregarious mountain ungulate (Alpine ibex, Capra ibex) over ten years focusing on groups, sub-groups and individuals, exploring the dynamics of sociality over different scales. Despite the social structure changing between seasons, the Alpine ibex population was highly cohesive: fission-fusion dynamics lead almost every male in the population to associate with each other male at least once. Nevertheless, we found that male Alpine ibex showed preferential associations that were maintained across seasons and years. Age seemed to be the most important factor driving preferential associations while other characteristics, such as social status, appeared less crucial. We also found that centrality measures were influenced by a...

Thermal imaging reveals social monitoring during social feeding in wild chimpanzees

Philosophical Transactions of the Royal Society B: Biological Sciences

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 s...

Male characteristics as predictors of genital color and display variation in vervet monkeys

Behavioral Ecology and Sociobiology, 2020

In the animal kingdom, conspicuous colors are often used for inter- and intra-sexual communication. Even though primates are the most colorful mammalian taxon, many questions, including what potential information color signals communicate to social partners, are not fully understood. Vervet monkeys (Chlorocebus pygerythrus) are ideal to examine the covariates of color signals. Males have multi-colored genitals, which they present during distinctive male-male interactions, known as the “Red-White-and-Blue” (RWB) display, but the genitals are also visible across a variety of other contexts, and it is unclear what this color display signals to recipients. We recorded genital color presentations and standardized digital photos of male genitals (N = 405 photos) over one mating season for 20 adult males in three groups at the Samara Private Game Reserve, South Africa. We combined these with data on male characteristics (dominance, age, tenure length, injuries, and fecal glucocorticoid met...

The prospect of rising in rank is key to long-term stability in Tibetan macaque society

Scientific reports, 2017

One of the most fundamental questions in behavioural biology is why societies can persist for a long period of time. While researchers in animal behaviour have been hindered by a lack of an aggregate measure (such as social mobility) to quantify the dynamics of animal societies, researchers in social sciences have been challenged by the complexity and diversity of human societies. As a result, direct empirical evidence is still lacking for the hypothesized causal relationship between social mobility and social stability. Here we attempt to fill the void by examining a much simpler society in the Tibetan macaque (Macaca thibetana), which we have tracked for 30 consecutive years. By testing two group-level hypotheses based on benefit-cost analysis and social stratification, we show the first quantitative evidence that an annual 2-to-1 stay/change ratio in the hierarchy with a 3-to-1 upward/downward ratio in intragenerational social mobility provides a substantive expected benefit for ...

The direct drivers of recent global anthropogenic biodiversity loss

Science Advances, 2022

Effective policies to halt biodiversity loss require knowing which anthropogenic drivers are the most important direct causes. Whereas previous knowledge has been limited in scope and rigor, here we statistically synthesize empirical comparisons of recent driver impacts found through a wide-ranging review. We show that land/sea use change has been the dominant direct driver of recent biodiversity loss worldwide. Direct exploitation of natural resources ranks second and pollution third; climate change and invasive alien species have been significantly less important than the top two drivers. The oceans, where direct exploitation and climate change dominate, have a different driver hierarchy from land and fresh water. It also varies among types of biodiversity indicators. For example, climate change is a more important driver of community composition change than of changes in species populations. Stopping global biodiversity loss requires policies and actions to tackle all the major d...

Chimpanzee fathers bias their behaviour towards their offspring

Royal Society open science, 2016

Promiscuous mating was traditionally thought to curtail paternal investment owing to the potential costs of providing care to unrelated infants. However, mounting evidence suggests that males in some promiscuous species can recognize offspring. In primates, evidence for paternal care exists in promiscuous Cercopithecines, but less is known about these patterns in other taxa. Here, we examine two hypotheses for paternal associations with lactating mothers in eastern chimpanzees (Pan troglodytes schweinfurthii): paternal effort, whereby males associate and interact more with their own infants, and mating effort, whereby males invest in mothers and offspring for mating privileges. We found that fathers associated more with their offspring than they did with non-kin infants, particularly early in life when infanticide risk is highest. Additionally, fathers and their infant offspring interacted more than expected. Notably, association between fathers and mother-infant pairs did not predi...

Group social rank is associated with performance on a spatial learning task

Royal Society open science, 2018

Dominant individuals differ from subordinates in their performances on cognitive tasks across a suite of taxa. Previous studies often only consider dyadic relationships, rather than the more ecologically relevant social hierarchies or networks, hence failing to account for how dyadic relationships may be adjusted within larger social groups. We used a novel statistical method: randomized Elo-ratings, to infer the social hierarchy of 18 male pheasants,, while in a captive, mixed-sex group with a linear hierarchy. We assayed individual learning performance of these males on a binary spatial discrimination task to investigate whether inter-individual variation in performance is associated with group social rank. Task performance improved with increasing trial number and was positively related to social rank, with higher ranking males showing greater levels of success. Motivation to participate in the task was not related to social rank or task performance, thus indicating that these ra...

Similar Task-Switching Performance of Real-Time Strategy and First-Person Shooter Players: Implications for Cognitive Training

Journal of Cognitive Enhancement, 2018

Computer games have been proposed as effective tools for cognitive enhancement. Especially first-person shooter (FPS) games have been found to yield a range of positive effects, and these positive effects also apply to the domain of executive functioning. Only a particular area of executive functioning has been shown to resist training via FPS games, and this area is task-switching performance. Here, we tested whether games of a different genre, real-time strategy (RTS) games, offer a more promising approach to improve task-switching performance, because RTS games capitalize on precisely this behavior. A high-powered, quasi-experimental comparison of RTS and FPS players indicated reliable costs for task-switching across both player groupswith similar performance on multiple indicators, comprising switch costs, mixing costs, voluntary switch rates, and psychological refractory period effects. Performance of both groups further did not exceed the performance of a control group of Chess and Go players. These results corroborate previous findings on the robustness of cognitive costs of task-switching. At the same time, our results also suggest that the precise characteristics of different computer games might not be critical in determining potential training effects. Keywords Task-switching. Executive functions. Computer games. Cognitive training. Cognitive enhancement BAy-yay-yay, the multi-tasking...the multi-tasking!M embTV, Caster (Age of Empires II)

Documenting Aggression, Dominance and the Impacts of Visitor Interaction on Galápagos Tortoises (Chelonoidis nigra) in a Zoo Setting

Animals, 2020

Ensuring high levels of welfare is imperative for modern zoos, but such organisations must also engage visitors in order to successfully spread awareness and raise conservation funds. It is therefore important to understand the responses of animals to visitor interaction to optimise welfare. Often, the opportunity to interact with humans may be enriching for animals, but in other contexts, this interaction may have negative welfare effects. We observed captive female Galápagos giant tortoises (Chelonoidis nigra) to describe aggressive interactions, characterize hierarchy using Elo ratings and assess the impact of visitor interactions. Elo ratings indicated that one individual was dominant over two equally ranked subordinates; aggressive interactions are discussed in this context. We detected significant effects of the presence of visitors and visitor type (keepers, vets or public) within the enclosure on aggression and activity. We suggest that previous miscategorisation of a natura...

Social relationships and greetings in wild chimpanzees (Pan troglodytes): use of signal combinations

Primates, 2019

Signals of submission, so-called 'greetings', represent an important tool for the regulation of social life in primates. In chimpanzees, vocalizations and gestures are commonly employed to communicate greetings, however, the topic of signal complexity (i.e., combinations of signals) during greeting instances has been neglected by research to date. Here, we investigate combinatorial possibilities in vocal greetings in a free-ranging group of chimpanzees (Pan troglodytes) and study how greeter sex, rank relationship between an interacting pair, and strength of the social bond of a greeting dyad influence signal complexity. Results show that the social bond and the dominance distance between individuals engaged in a greeting bout are important determiners for vocal combinations. The findings indicate that greeting signals in chimpanzees, like other vocal signals of the species, can become subject to social influences.

Climate, Temporal Abundance of Key Food Sources and Home Ranges of Crested Macaques (Macaca nigra) in Sulawesi, Indonesia: A Longitudinal Phenological Study

International Journal of Primatology

Climate change is associated with more frequent extreme weather conditions and an overall increase in temperature around the globe. Its impact on individual ecosystems is not yet well known. Long-term data documenting climate and the temporal abundance of food for primates are scarce. We used long-term phenological data to assess climate variation, fruit abundance and home range sizes of the endemic and Critically Endangered crested macaques (Macaca nigra) in Tangkoko forest, Sulawesi, Indonesia. Between January 2012 and July 2020, every month, we monitored 498 individual trees from 41 species and 23 families. We noted each tree’s phenophase and assessed variation in climate (daily temperature and rainfall) and fruit abundance. We also investigated whether individual trees of known key food sources for macaques (New Guinea walnut trees, Dracontomelon spp, two species, N = 10 individual trees; fig trees, Ficus spp, four species, N = 34, and spiked peppers, Piper aduncum, N = 4) showe...

Considering Social Play in Primates: A Case Study in Juvenile Tibetan Macaques (Macaca thibetana)

The Behavioral Ecology of the Tibetan Macaque

Social play in animals has long intrigued ethologists, and current research in this field, especially pertaining to primates, has revealed that the behavioral lives of juveniles in particular are rich and behaviorally and socially complex. It is widely acknowledged that play contributes to the biological, social, and cognitive growth of juveniles, but the impact of this growth on fitness, future adult social relationships, or general social group experiences remains unknown. Comparative social play research can enhance our understanding of how play influences these variables critical to juvenile development and fitness. Comparisons across taxa with varying social organizations, e.g., Macaca spp., can further this understanding. In this chapter, we introduce the 2017 social play network of Yulingkeng A1 infant and juvenile Tibetan macaques (Macaca thibetana) and discuss how social play is organized in this species.

Discovering Implied Serial Order Through Model-Free and Model-Based Learning

Frontiers in Neuroscience, 2019

Humans and animals can learn to order a list of items without relying on explicit spatial or temporal cues. To do so, they appear to make use of transitivity, a property of all ordered sets. Here, we summarize relevant research on the transitive inference (TI) paradigm and its relationship to learning the underlying order of an arbitrary set of items. We compare six computational models of TI performance, three of which are modelfree (Q-learning, Value Transfer, and REMERGE) and three of which are model-based (RL-Elo, Sequential Monte Carlo, and Betasort). Our goal is to assess the ability of these models to produce empirically observed features of TI behavior. Model-based approaches perform better under a wider range of scenarios, but no single model explains the full scope of behaviors reported in the TI literature.

Juvenile rank acquisition is associated with fitness independent of adult rank

Proceedings of the Royal Society B: Biological Sciences, 2020

Social rank is a significant determinant of fitness in a variety of species. The importance of social rank suggests that the process by which juveniles come to establish their position in the social hierarchy is a critical component of development. Here, we use the highly predictable process of rank acquisition in spotted hyenas to study the consequences of variation in rank acquisition in early life. In spotted hyenas, rank is ‘inherited’ through a learning process called ‘maternal rank inheritance.’ This pattern is very consistent: approximately 80% of juveniles acquire the exact rank expected under the rules of maternal rank inheritance. The predictable nature of rank acquisition in these societies allows the process of rank acquisition to be studied independently from the ultimate rank that each juvenile attains. In this study, we use Elo-deviance scores, a novel application of the Elo-rating method, to calculate each juvenile's deviation from the expected pattern of materna...

Trade-offs in the production of animal vocal sequences: insights from the structure of wild chimpanzee pant hoots

Frontiers in Zoology, 2017

Background: Vocal sequences-utterances consisting of calls produced in close succession-are common phenomena in animal communication. While many studies have explored the adaptive benefits of producing such sequences, very little is known about how the costs and constraints involved in their production affect their form. Here, we investigated this issue in the chimpanzee (Pan troglodytes schweinfurthii) pant hoot, a long and structurally complex vocal sequence comprising four acoustically distinct phasesintroduction, build-up, climax and let-down. Results: We found that in each of these phases, and for the sequence as a whole, there was a negative relationship between the number of calls produced and their average duration. There was also a negative relationship between the total duration of some adjacent phases. Significant relationships between the fundamental frequency of calls and their number or duration were found for some phases of the sequence, but the direction of these relationships differed between particular phases. Conclusions: These results indicate that there are trade-offs in terms of signal duration at two levels in pant-hoot production: between call number and duration, and between the relative durations of successive phases. These trade-offs are likely to reflect biomechanical constraints on vocal sequence production. Phase-specific trade-offs also appear to occur between fundamental frequency and call number or duration, potentially reflecting that different phases of the sequence are associated with distinct types of information, linked in different ways to call pitch. Overall, this study highlights the important role of costs and constraints in shaping the temporal and acoustic structure of animal vocal sequences.

A first exploratory comparison of the behaviour of wolves (Canis lupus) and wolf-dog hybrids in captivity

Research Square (Research Square), 2023

Extensive introgression of genes from domesticated taxa may be a serious threat for the genomic integrity and adaptability of wild populations. Grey wolves (Canis lupus) are especially vulnerable to this phenomenon, but there are no studies yet assessing the potential behavioural effects of dogintrogression in wolves. In this study, we conducted a rst systematic comparison of admixed (N = 11) and non-admixed wolves (N = 14) in captivity, focusing on their reaction to unfamiliar humans and novel objects, and the cohesiveness of their social groups. When exposed to unfamiliar humans in the experimental task, wolves were more vigilant, fearful and aggressive than admixed wolves, and less likely to approach humans, but also more likely to spend time in human proximity. When exposed to novel objects, wolves were more aggressive than admixed wolves, less likely to spend time in object proximity, and more likely to interact with objects, but also less vigilant and as fearful as admixed wolves. Finally, social networks were mostly more cohesive in wolves than admixed wolves. Overall, our study suggests that dog admixture may lead to important behavioural changes in wolves, with possible implications for conservation strategies.

Measuring dominance certainty and assessing its impact on individual and societal health in a nonhuman primate model: a network approach

Philosophical Transactions of the Royal Society B: Biological Sciences, 2022

The notion of dominance is ubiquitous across the animal kingdom, wherein some species/groups such relationships are strictly hierarchical and others are not. Modern approaches for measuring dominance have emerged in recent years taking advantage of increased computational power. One such technique, named Percolation and Conductance (Perc), uses both direct and indirect information about the flow of dominance relationships to generate hierarchical rank order that makes no assumptions about the linearity of these relationships. It also provides a new metric, known as ‘dominance certainty’, which is a complimentary measure to dominance rank that assesses the degree of ambiguity of rank relationships at the individual, dyadic and group levels. In this focused review, we will (i) describe how Perc measures dominance rank while accounting for both nonlinear hierarchical structure as well as sparsity in data—here we also provide a metric of dominance certainty estimated by Perc, which can ...

Inferring longitudinal hierarchies: Framework and methods for studying the dynamics of dominance

Journal of Animal Ecology, 2019

1. Social inequality is a consistent feature of animal societies, often manifesting as dominance hierarchies, in which each individual is characterized by a dominance rank denoting its place in the network of competitive relationships among group members. Most studies treat dominance hierarchies as static entities despite their true longitudinal, and sometimes highly dynamic, nature.

The function of chimpanzee greeting calls is modulated by their acoustic variation

Animal Behaviour, 2021

Signalling plays an important role in mediating social interactions in many animal species. For example, during approaches certain species produce "greetings", which can take the form of vocal or visual signals, which reduce the probability of aggressive interactions and/or facilitate affiliation when approaching each other. However, in species where greetings are comprised of both vocal and visual signals, little is known about how the vocal component relates to the visual component, or, in species with fission-fusion dynamics, to the time spent together by the dyad in the same subgroup prior to the approach. Similarly, in species with several acoustic variants of greeting calls, it is unclear whether different variants have different functions. We looked at the production of two acoustically distinct greeting call variants, low-fundamental frequency pant grunts and high-fundamental frequency pant barks, during approaches between two individuals in five communities of wild chimpanzees (Pan troglodytes) in Uganda and Ivory Coast. More specifically, we explored the relationship between greeting call production and i) aggressive and submissive interactions during the approach ii) preceding and subsequent proximity levels between the involved individuals. Calls were more likely to be produced during aggressive interactions and were associated with postures and gestures linked to submission; these patterns were stronger when the utterance contained a pant bark rather than a pant grunt alone. The production of greeting calls was more likely soon after party fusion and was negatively related to subsequent proximity levels between the caller and receiver. These results expand our knowledge of greeting calls and imply that these calls might be used to re-establish dominance relationships after a period of separation, and that the function of these calls can be modulated by their specific acoustic variants and by visual signals that often accompany them.

An intentional cohesion call in male chimpanzees of Budongo Forest

Animal Cognition, 2022

Many social animals travel in cohesive groups but some species, including chimpanzees, form flexible fission-fusion systems where individuals have some control over group cohesion and proximity to others. Here, we explored how male chimpanzees of the Sonso community of Budongo Forest, Uganda, use communication signals during resting, a context where the likelihood of group fission is high due to forthcoming travel. We focused on a context-specific vocalisation, the 'rest hoo', to investigate its function and determine whether it is produced intentionally. We found that this call was typically given towards the end of typical silent resting bouts, i.e., the period when individuals need to decide whether to continue travelling after a brief stop-over or to start a prolonged resting bout. Subjects rested longer after producing 'rest hoos' and their resting time increased with the number of calls produced. They also rested longer if their calls were answered. Furthermore, focal subjects' resting time was prolonged after hearing others' 'rest hoos'. Subjects called more when with top proximity partners and in small parties and rested longer if a top proximity partner called. We also found an interaction effect between rank and grooming activity, with high-ranking males with a high grooming index calling less frequently than other males, suggesting that vocal communication may serve as a cohesion strategy alternative to tactile-based bonding. We discuss these different patterns and conclude that chimpanzee 'rest hoos' meet key criteria for intentional signalling. We suggest that 'rest hoos' are produced to prolong resting bouts with desired partners, which may function to increase social cohesion.

The Evolution of Food Calls: Vocal Behaviour of Sooty Mangabeys in the Presence of Food

Frontiers in Psychology, 2022

The two main theories of food-associated calls in animals propose functions either in cooperative recruitment or competitive spacing. However, not all social animals produce food calls and it is largely unclear under what circumstances this call type evolves. Sooty mangabeys (Cercocebus atys) do not have food calls, but they frequently produce grunts during foraging, their most common vocalisation. We found that grunt rates were significantly higher when subjects were foraging in the group's periphery and with small audiences, in line with the cooperative recruitment hypothesis. In a subsequent field experiment we presented highly desired food items and found that discovering individuals called, unless harassed by competitors, but that the calls never attracted others, confirming that the grunts do not convey any information referential to food. Our data thus suggest that the evolution of cooperative food calling is a two-step process, starting with increased motivation to vocalise in the feeding context, followed by the evolution of acoustic variants derived from context-general contact calls. This evolutionary transition may only occur in species that feed on clumped, high-quality resources where social feeding is competitive, a condition not met in sooty mangabeys.

Risk factors for respiratory illness in a community of wild chimpanzees ( Pan troglodytes schweinfurthii )

Royal Society Open Science, 2018

Respiratory illnesses have caused significant mortality in African great ape populations. While much effort has been given to identifying the responsible pathogens, little is known about the factors that influence disease transmission or individual susceptibility. In the Kanyawara community of wild chimpanzees, respiratory illness has been the leading cause of mortality over 31 years, contributing to 27% of deaths. Deaths were common in all age groups except juveniles. Over 22 years of health observations, respiratory signs were rare among infants and most common among older adults of both sexes. Respiratory signs were also common among males during the transition to adulthood (ages 10-20 years), particularly among those of low rank. Respiratory signs peaked conspicuously in March, a pattern that we could not explain after modelling climatic factors, group sizes, diet or exposure to humans. Furthermore, rates of respiratory illness in the chimpanzees did not track seasonal rates of illness in the nearby village. Our data indicate that the epidemiology of chimpanzee respiratory illness warrants more investigation but clearly differs in important ways from humans. Findings on individual susceptibility patterns suggest that respiratory signs are a robust indicator for investigating immunocompetence in wild chimpanzees.

Greeting Behaviors in Male Alouatta palliata at La Pacifica, Costa Rica

International Journal of Primatology, 2019

Greeting behaviors are ritualized, nonaggressive interactions that serve as a form of tactile communication between two individuals; however, our understanding of the function of male greetings and how they vary inter-and intraspecifically is limited, particularly in Neotropical species. I examined greeting behavior in male Alouatta palliata to determine their social function by testing the tension reduction, social bonding, and formal dominance hypotheses. I collected 1751 h of behavioral data on two groups (G2 and G12/42) of similar size and sex ratio, with clear male dominance hierarchies. The overall number of greetings was low in both groups (N = 36 for both groups). More greetings occurred during times of social disruption (i.e., intergroup encounters, competition for food or females) than during neutral contexts (i.e., traveling or resting without potentially cycling females nearby), though frequencies varied by context in each group. Greetings occurred during all intergroup encounters in G2 but not in G12/42. With the exception of the competitive context in G12/42, alpha males and the higher-ranked male in the dyad initiated the majority of greetings in all contexts, though unidirectionality was not consistent. The pattern of greetings differed among dyads and between the groups. Subordinates in the dyad initiated more greetings than dominants only in G12/42 in the context of competitive activities, and the number of greetings decreased in G12/42 as rank distance increased. These results partially support the tension reduction hypothesis, and suggest greetings communicate nonagonistic intent among males. However, the nature of these encounters can vary by dyad and group within the same population, suggesting that greetings may also be partly driven by differing group histories, the social and ecological environments surrounding each group, and individual male temperaments.