Single aggressive interactions increase urinary glucocorticoid levels in wild male chimpanzees

Financial support was provided by British Academy [http://www.britac.ac.uk/] (CC), Leakey Foundation [http://www.leakeyfoundation.org/] (CC, RMW, TD), Leverhulm Trust [http://www.leverhulme.ac.uk/] (KZ), Max Planck Society [http://www.eva.mpg.de/] (AW, CC, RMW, TD) and the Royal Zoological Society of Scotland [http://www.rzss.org.uk/], in providing core funding for Budongo Conservation Field Station. This project has received additional funding from the European Union’s Seventh Framework Program for research, technological development and demonstration [http://ec.europa.eu/research/fp7/index_e​n.cfm] under grant agreement no 283871.A basic premise in behavioural ecology is the cost-benefit arithmetic, which determines both behavioural decisions and evolutionary processes. Aggressive interactions can be costly on an energetic level, demanding increased energy or causing injuries, and on a psychological level, in the form of increased anxiety and damaged relationships between opponents. Here we used urinary glucocorticoid (uGC) levels to assess the costs of aggression in wild chimpanzees of Budongo Forest, Uganda. We collected 169 urine samples from nine adult male chimpanzees following 14 aggressive interactions (test condition) and 10 resting events (control condition). Subjects showed significantly higher uGC levels after single aggressive interactions compared to control conditions, likely for aggressors as well as victims. Higher ranking males had greater increases of uGC levels after aggression than lower ranking males. In contrast, uGC levels showed no significant change in relation to aggression length or intensity, indicating that psychological factors might have played a larger role than mere energetic expenditure. We concluded that aggressive behaviour is costly for both aggressors and victims and that costs seem poorly explained by energetic demands of the interaction. Our findings are relevant for studies of post-conflict interactions, since we provide evidence that both aggressors and victims experience a stress response to conflict.Publisher PDFPeer reviewe

The syntax and meaning of wild gibbon songs

Spoken language is a result of the human capacity to assemble simple vocal units into more complex utterances, the basic carriers of semantic information. Not much is known about the evolutionary origins of this behaviour. The vocal abilities of non-human primates are relatively unimpressive in comparison, with gibbon songs being a rare exception. These apes assemble a repertoire of call notes into elaborate songs, which function to repel conspecific intruders, advertise pair bonds, and attract mates. We conducted a series of field experiments with white-handed gibbons at Khao Yai National Park, Thailand, which showed that this ape species uses songs also to protect themselves against predation. We compared the acoustic structure of predatory-induced songs with regular songs that were given as part of their daily routine. Predator-induced songs were identical to normal songs in the call note repertoire, but we found consistent differences in how the notes were assembled into songs. The responses of out-of-sight receivers demonstrated that these syntactic differences were meaningful to conspecifics. Our study provides the first evidence of referential signalling in a free-ranging ape species, based on a communication system that utilises combinatorial rules

Vocal communication in gibbons.

Many non-human primates use vocal communication referentially and also use simple syntax and grammar. However, their comparative vocal repertoires are disappointingly sparse, with many researchers concluding that they have fixed vocal patterns made up of a limited number of discrete units used in a relatively small array of contexts (see McComb & Semple, 2005 for a review). Furthermore, these vocal patterns seem to be innate, under high genetic control with little evidence for vocal learning – something that humans are masters at (Janik & Slater 1997). This leaves us with some questions. Firstly, how did humans become so adept at producing and learning vocal sounds? And, secondly, are there any extant primate species with vocal behaviours that can be directly compared to our own?

Juvenile vervet monkeys rely on others when responding to danger

Funding: Open access funding provided by University of Lausanne. Funding was provided by FP7 Ideas: European Research Council, 283871, Schweizerischer Nationalfonds zur Förderung der Wissenschaftlichen Forschung, 310030_143359, PP00P3_170624, Branco Weiss Fellowship-Society in Science.Primate alarm calls are mainly hardwired but individuals need to adapt their calling behaviours according to the situation. Such learning necessitates recognising locally relevant dangers and may take place via their own experience or by observing others. To investigate monkeys alarm calling behaviour, we carried out a field experiment in which we exposed juvenile vervet monkeys to unfamiliar raptor models in the presence of audiences that differed in experience and reliability. We used audience age as a proxy for experience and relatedness as a proxy for reliability, while quantifying audience reactions to the models. We found a negative correlation between alarm call production and callers’ age. Adults never alarm called, compared to juveniles. We found no overall effect of audience composition and size, with juveniles calling more when with siblings than mothers or unrelated individuals. Finally, concerning audience reactions to the models, we observed juveniles remained silent with vigilant mothers and only alarm called with ignoring mothers, whereas we observed the opposite for siblings: juveniles remained silent with ignoring siblings and called with vigilant siblings. Despite the small sample size, juvenile vervet monkeys, confronted with unfamiliar and potentially dangerous raptors, seem to rely on others to decide whether to alarm call, demonstrating that the choice of the model may play an important key role in the ontogeny of primate alarm call behaviour.Publisher PDFPeer reviewe

Vocal communication in gibbons

Many non-human primates use vocal communication referentially and also use simple syntax and grammar. However, their comparative vocal repertoires are disappointingly sparse, with many researchers concluding that they have fixed vocal patterns made up of a limited number of discrete units used in a relatively small array of contexts (see McComb & Semple, 2005 for a review). Furthermore, these vocal patterns seem to be innate, under high genetic control with little evidence for vocal learning – something that humans are masters at (Janik & Slater 1997). This leaves us with some questions. Firstly, how did humans become so adept at producing and learning vocal sounds? And, secondly, are there any extant primate species with vocal behaviours that can be directly compared to our own?

Evidence of joint commitment in great apes’ natural joint actions

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 we discuss possible evolutionary origins

Campbell's Monkeys Use Affixation to Alter Call Meaning

Human language has evolved on a biological substrate with phylogenetic roots deep in the primate lineage. Here, we describe a functional analogy to a common morphological process in human speech, affixation, in the alarm calls of free-ranging adult Campbell's monkeys (Cercopithecus campbelli campbelli). We found that male alarm calls are composed of an acoustically variable stem, which can be followed by an acoustically invariable suffix. Using long-term observations and predator simulation experiments, we show that suffixation in this species functions to broaden the calls' meaning by transforming a highly specific eagle alarm to a general arboreal disturbance call or by transforming a highly specific leopard alarm call to a general alert call. We concluded that, when referring to specific external events, non-human primates can generate meaningful acoustic variation during call production that is functionally equivalent to suffixation in human language

Correlates of social role and conflict severity in wild vervet monkey agonistic screams.

Screams are acoustically distinct, high-pitched and high-amplitude calls, produced by many social species. Despite a wide range of production contexts, screams are characterised by an acoustic structure that appears to serve in altering the behaviour of targeted receivers during agonistic encounters. In chimpanzees, this can be achieved by callers producing acoustic variants that correlate with their identity, social role, relationship with the targeted recipient, the composition of the audience and the nature of the event. Although vervet monkeys (Chlorocebus pygerythrus) have been studied for decades, not much is known about their agonistic screams. Here, we examined agonistic screams produced by wild vervet monkeys to investigate the degree to which caller identity, social role and conflict severity affected call structure. We found that screams were both individually distinctive and dependent of the agonistic events. In particular, victim screams were longer and higher-pitched than aggressor screams, while screams produced in severe conflicts (chases, physical contact) had higher entropy than those in mild conflicts. We discuss these findings in terms of their evolutionary significance and suggest that acoustic variation might serve to reduce the aggression level of opponents, while simultaneously attracting potential helpers

Quantifying uncertainty due to fission-fusion dynamics as a component of social complexity.

Groups of animals (including humans) may show flexible grouping patterns, in which temporary aggregations or subgroups come together and split, changing composition over short temporal scales, (i.e. fission and fusion). A high degree of fission-fusion dynamics may constrain the regulation of social relationships, introducing uncertainty in interactions between group members. Here we use Shannon's entropy to quantify the predictability of subgroup composition for three species known to differ in the way their subgroups come together and split over time: spider monkeys (Ateles geoffroyi), chimpanzees (Pan troglodytes) and geladas (Theropithecus gelada). We formulate a random expectation of entropy that considers subgroup size variation and sample size, against which the observed entropy in subgroup composition can be compared. Using the theory of set partitioning, we also develop a method to estimate the number of subgroups that the group is likely to be divided into, based on the composition and size of single focal subgroups. Our results indicate that Shannon's entropy and the estimated number of subgroups present at a given time provide quantitative metrics of uncertainty in the social environment (within which social relationships must be regulated) for groups with different degrees of fission-fusion dynamics. These metrics also represent an indirect quantification of the cognitive challenges posed by socially dynamic environments. Overall, our novel methodological approach provides new insight for understanding the evolution of social complexity and the mechanisms to cope with the uncertainty that results from fission-fusion dynamics

Chimpanzees Extract Social Information from Agonistic Screams

Chimpanzee (Pan troglodytes) agonistic screams are graded vocal signals that are produced in a context-specific manner. Screams given by aggressors and victims can be discriminated based on their acoustic structure but the mechanisms of listener comprehension of these calls are currently unknown. In this study, we show that chimpanzees extract social information from these vocal signals that, combined with their more general social knowledge, enables them to understand the nature of out-of-sight social interactions. In playback experiments, we broadcast congruent and incongruent sequences of agonistic calls and monitored the response of bystanders. Congruent sequences were in accordance with existing social dominance relations; incongruent ones violated them. Subjects looked significantly longer at incongruent sequences, despite them being acoustically less salient (fewer call types from fewer individuals) than congruent ones. We concluded that chimpanzees categorised an apparently simple acoustic signal into victim and aggressor screams and used pragmatics to form inferences about third-party interactions they could not see