Animal communication has long been thought to be subject to pressures and constraints associated ... more Animal communication has long been thought to be subject to pressures and constraints associated with social relationships. However, our understanding of how the nature and quality of social relationships relates to the use and evolution of communication is limited by a lack of directly comparable methods across multiple levels of analysis. Here, we analysed observational data from 111 wild groups belonging to 26 non-human primate species, to test how vocal communication relates to dominance style (the strictness with which a dominance hierarchy is enforced, ranging from 'despotic' to 'tolerant'). At the individual-level, we found that dominant individuals who were more tolerant vocalized at a higher rate than their despotic counterparts. This indicates that tolerance within a relationship may place pressure on the dominant partner to communicate more during social interactions. At the species-level, however, despotic species exhibited a larger repertoire of hierarch...
This is the csv data file including all individual level data; suitable for analyses in R script ... more This is the csv data file including all individual level data; suitable for analyses in R script entitled 'Individual_Level_Models'
This is the csv data file including all species level data; suitable for analyses using R script ... more This is the csv data file including all species level data; suitable for analyses using R script entitled 'Species_Level_Models'
Animal communication has long been thought to be subject to pressures and constraints associated ... more Animal communication has long been thought to be subject to pressures and constraints associated with social relationships. However, our understanding of how the nature and quality of social relationships relates to the use and evolution of communication is limited by a lack of directly comparable methods across multiple levels of analysis. Here, we analysed observational data from 111 wild groups belonging to 26 non-human primate species, to test how vocal communication relates to dominance style (the strictness with which a dominance hierarchy is enforced, ranging from ‘despotic’ to ‘tolerant’). At the individual-level, we found that dominant individuals who were more tolerant vocalized at a higher rate than their despotic counterparts. This indicates that tolerance within a relationship may place pressure on the dominant partner to communicate more during social interactions. At the species-level, however, despotic species exhibited a larger repertoire of hierarchy-related vocali...
Animals communicate using a vast array of different signals in different modalities. For chimpanz... more Animals communicate using a vast array of different signals in different modalities. For chimpanzees, vocalizations, gestures and facial expressions are all important forms of communication, yet these signals have rarely been studied together holistically. The current study aimed to provide the first comprehensive repertoire of flexibly combined ('free') multimodal (MM) signals, and assess individual and contextual factors influencing production of, and responses to, unimodal (UM) and MM signals in wild chimpanzees. In total, 48 different free MM signals were produced. MM signals were produced at a significantly lower rate than UM signals, but 22 of 26 focal animals were observed to produce free MM signals. The relative production rates of different types of UM and MM signals differed significantly between the behavioural contexts investigated, showing flexible use of signals across contexts. In contrast, individual factors such as age, sex or rank of signaller did not appear to influence the type of signal produced or the likelihood of eliciting a response. Finally, we compared recipient responses to free MM grunt-gesture signals and matched UM component signals and found that these MM signals were more likely to elicit a response than a grunt alone, but were as likely to elicit a response as the gesture alone. The overall findings point to a widespread capacity for wild chimpanzees to flexibly combine signals from different modalities and highlight the importance of adopting a multimodal approach to studying communication.
Biological reviews of the Cambridge Philosophical Society, Jan 2, 2016
Language's intentional nature has been highlighted as a crucial feature distinguishing it fro... more Language's intentional nature has been highlighted as a crucial feature distinguishing it from other communication systems. Specifically, language is often thought to depend on highly structured intentional action and mutual mindreading by a communicator and recipient. Whilst similar abilities in animals can shed light on the evolution of intentionality, they remain challenging to detect unambiguously. We revisit animal intentional communication and suggest that progress in identifying analogous capacities has been complicated by (i) the assumption that intentional (that is, voluntary) production of communicative acts requires mental-state attribution, and (ii) variation in approaches investigating communication across sensory modalities. To move forward, we argue that a framework fusing research across modalities and species is required. We structure intentional communication into a series of requirements, each of which can be operationalised, investigated empirically, and must...
Long-distance calling is a common behaviour in animals that has various important social function... more Long-distance calling is a common behaviour in animals that has various important social functions. At a physiological level, calling is often mediated by gonadal hormones such as testosterone (T), particularly when its function is linked to intra-sexual competition for mates or territory. T also plays an important role in the development of vocal characteristics associated with dominance in humans. However, the few available studies of T and vocal behaviour in non-human primates suggest that in primates T has less influence on call production than in other animals. We tested this hypothesis by studying the relationship between T concentrations and pant hooting in wild male chimpanzees (Pan troglodytes schweinfurthii) of the Kanyawara community in the Kibale National Park, Uganda. We found three kinds of correlation. Hourly T averages were positively associated with hourly rates of pant-hooting. Monthly T levels were likewise correlated with monthly rates of pant hooting after controlling for other influences such as fission-fusion rates. Finally, males with high T levels had higher peak frequency at the start of the call climax. These results suggest that T affects the production of pant-hoots in chimpanzees. This implies that the pant-hoot call plays a role in male-male competition. We propose that even in cognitively sophisticated species, endocrine mechanisms can contribute to regulating vocal production.
BioOne Complete (complete.BioOne.org) is a full-text database of 200 subscribed and open-access t... more BioOne Complete (complete.BioOne.org) is a full-text database of 200 subscribed and open-access titles in the biological, ecological, and environmental sciences published by nonprofit societies, associations, museums, institutions, and presses.
Scientists studying the communication of non-human animals are often aiming to better understand ... more Scientists studying the communication of non-human animals are often aiming to better understand the evolution of human communication, including human language. Some scientists take a phylogenetic perspective, where the goal is to trace the evolutionary history of communicative traits, while others take a functional perspective, where the goal is to understand the selection pressures underpinning specific traits. Both perspectives are necessary to fully understand the evolution of communication, but it is important to understand how the two perspectives differ and what they can and cannot tell us. Here, we suggest that integrating phylogenetic and functional questions can be fruitful in better understanding the evolution of communication. We also suggest that adopting a multimodal approach to communication might help to integrate phylogenetic and functional questions, and provide an interesting avenue for research into language evolution.
Signalling plays an important role in facilitating and maintaining affiliative or cooperative int... more 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...
Long-distance calls have a variety of functions in different animal species. However, where multi... more Long-distance calls have a variety of functions in different animal species. However, where multiple functions are proposed for a single long-distance call type, little is known about their relative importance. Chimpanzees are one species where several functions have been proposed for their long-distance call, the pant hoot. In this study, we investigated the effect of social factors, including the rank of the caller, party size, fission-fusion rates, and the presence of estrus females as well as ecological factors including the type of food consumed and travel time, on male chimpanzee (Pan troglodytes schweinfurthii) pant hooting, in order to identify the key correlates of this behavior. The wild male chimpanzees of the Kanyawara community, Uganda, produced more pant hoots on days when there were frequent changes in the male, but not female, composition of the focal's party. This factor accounted for the largest amount of variation in pant hoot production, and we found that males were more likely to repeat a call prior to rather than after fusion with other males, suggesting that the calls facilitate fusion. Pant hoots therefore seem to play a pivotal role in regulating grouping dynamics in chimpanzees. Our study also shows that pant hooting was positively correlated with the rank of the caller, the presence of parous females in estrus, and the consumption of highquality food, suggesting that pant hoots signal social status or social bonds when between-male competition is high. This study supports the view that pant hoots fulfill a complex social function.
The production of structured and repetitive sounds by striking objects is a behavior found not on... more The production of structured and repetitive sounds by striking objects is a behavior found not only in humans, but also in a variety of animal species, including chimpanzees (Pan troglodytes). In this study we examined individual and social factors that may influence the frequency with which individuals engage in drumming behavior when producing long distance pant hoot vocalizations, and analyzed the temporal structure of those drumming bouts. Male chimpanzees from Budongo Forest, Uganda, drummed significantly more frequently during travel than feeding or resting and older individuals were significantly more likely to produce drumming bouts than younger ones. In contrast, we found no evidence that the presence of estrus females, high ranking males and
Animal communication has long been thought to be subject to pressures and constraints associated ... more Animal communication has long been thought to be subject to pressures and constraints associated with social relationships. However, our understanding of how the nature and quality of social relationships relates to the use and evolution of communication is limited by a lack of directly comparable methods across multiple levels of analysis. Here, we analysed observational data from 111 wild groups belonging to 26 non-human primate species, to test how vocal communication relates to dominance style (the strictness with which a dominance hierarchy is enforced, ranging from 'despotic' to 'tolerant'). At the individual-level, we found that dominant individuals who were more tolerant vocalized at a higher rate than their despotic counterparts. This indicates that tolerance within a relationship may place pressure on the dominant partner to communicate more during social interactions. At the species-level, however, despotic species exhibited a larger repertoire of hierarch...
This is the csv data file including all individual level data; suitable for analyses in R script ... more This is the csv data file including all individual level data; suitable for analyses in R script entitled 'Individual_Level_Models'
This is the csv data file including all species level data; suitable for analyses using R script ... more This is the csv data file including all species level data; suitable for analyses using R script entitled 'Species_Level_Models'
Animal communication has long been thought to be subject to pressures and constraints associated ... more Animal communication has long been thought to be subject to pressures and constraints associated with social relationships. However, our understanding of how the nature and quality of social relationships relates to the use and evolution of communication is limited by a lack of directly comparable methods across multiple levels of analysis. Here, we analysed observational data from 111 wild groups belonging to 26 non-human primate species, to test how vocal communication relates to dominance style (the strictness with which a dominance hierarchy is enforced, ranging from ‘despotic’ to ‘tolerant’). At the individual-level, we found that dominant individuals who were more tolerant vocalized at a higher rate than their despotic counterparts. This indicates that tolerance within a relationship may place pressure on the dominant partner to communicate more during social interactions. At the species-level, however, despotic species exhibited a larger repertoire of hierarchy-related vocali...
Animals communicate using a vast array of different signals in different modalities. For chimpanz... more Animals communicate using a vast array of different signals in different modalities. For chimpanzees, vocalizations, gestures and facial expressions are all important forms of communication, yet these signals have rarely been studied together holistically. The current study aimed to provide the first comprehensive repertoire of flexibly combined ('free') multimodal (MM) signals, and assess individual and contextual factors influencing production of, and responses to, unimodal (UM) and MM signals in wild chimpanzees. In total, 48 different free MM signals were produced. MM signals were produced at a significantly lower rate than UM signals, but 22 of 26 focal animals were observed to produce free MM signals. The relative production rates of different types of UM and MM signals differed significantly between the behavioural contexts investigated, showing flexible use of signals across contexts. In contrast, individual factors such as age, sex or rank of signaller did not appear to influence the type of signal produced or the likelihood of eliciting a response. Finally, we compared recipient responses to free MM grunt-gesture signals and matched UM component signals and found that these MM signals were more likely to elicit a response than a grunt alone, but were as likely to elicit a response as the gesture alone. The overall findings point to a widespread capacity for wild chimpanzees to flexibly combine signals from different modalities and highlight the importance of adopting a multimodal approach to studying communication.
Biological reviews of the Cambridge Philosophical Society, Jan 2, 2016
Language's intentional nature has been highlighted as a crucial feature distinguishing it fro... more Language's intentional nature has been highlighted as a crucial feature distinguishing it from other communication systems. Specifically, language is often thought to depend on highly structured intentional action and mutual mindreading by a communicator and recipient. Whilst similar abilities in animals can shed light on the evolution of intentionality, they remain challenging to detect unambiguously. We revisit animal intentional communication and suggest that progress in identifying analogous capacities has been complicated by (i) the assumption that intentional (that is, voluntary) production of communicative acts requires mental-state attribution, and (ii) variation in approaches investigating communication across sensory modalities. To move forward, we argue that a framework fusing research across modalities and species is required. We structure intentional communication into a series of requirements, each of which can be operationalised, investigated empirically, and must...
Long-distance calling is a common behaviour in animals that has various important social function... more Long-distance calling is a common behaviour in animals that has various important social functions. At a physiological level, calling is often mediated by gonadal hormones such as testosterone (T), particularly when its function is linked to intra-sexual competition for mates or territory. T also plays an important role in the development of vocal characteristics associated with dominance in humans. However, the few available studies of T and vocal behaviour in non-human primates suggest that in primates T has less influence on call production than in other animals. We tested this hypothesis by studying the relationship between T concentrations and pant hooting in wild male chimpanzees (Pan troglodytes schweinfurthii) of the Kanyawara community in the Kibale National Park, Uganda. We found three kinds of correlation. Hourly T averages were positively associated with hourly rates of pant-hooting. Monthly T levels were likewise correlated with monthly rates of pant hooting after controlling for other influences such as fission-fusion rates. Finally, males with high T levels had higher peak frequency at the start of the call climax. These results suggest that T affects the production of pant-hoots in chimpanzees. This implies that the pant-hoot call plays a role in male-male competition. We propose that even in cognitively sophisticated species, endocrine mechanisms can contribute to regulating vocal production.
BioOne Complete (complete.BioOne.org) is a full-text database of 200 subscribed and open-access t... more BioOne Complete (complete.BioOne.org) is a full-text database of 200 subscribed and open-access titles in the biological, ecological, and environmental sciences published by nonprofit societies, associations, museums, institutions, and presses.
Scientists studying the communication of non-human animals are often aiming to better understand ... more Scientists studying the communication of non-human animals are often aiming to better understand the evolution of human communication, including human language. Some scientists take a phylogenetic perspective, where the goal is to trace the evolutionary history of communicative traits, while others take a functional perspective, where the goal is to understand the selection pressures underpinning specific traits. Both perspectives are necessary to fully understand the evolution of communication, but it is important to understand how the two perspectives differ and what they can and cannot tell us. Here, we suggest that integrating phylogenetic and functional questions can be fruitful in better understanding the evolution of communication. We also suggest that adopting a multimodal approach to communication might help to integrate phylogenetic and functional questions, and provide an interesting avenue for research into language evolution.
Signalling plays an important role in facilitating and maintaining affiliative or cooperative int... more 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...
Long-distance calls have a variety of functions in different animal species. However, where multi... more Long-distance calls have a variety of functions in different animal species. However, where multiple functions are proposed for a single long-distance call type, little is known about their relative importance. Chimpanzees are one species where several functions have been proposed for their long-distance call, the pant hoot. In this study, we investigated the effect of social factors, including the rank of the caller, party size, fission-fusion rates, and the presence of estrus females as well as ecological factors including the type of food consumed and travel time, on male chimpanzee (Pan troglodytes schweinfurthii) pant hooting, in order to identify the key correlates of this behavior. The wild male chimpanzees of the Kanyawara community, Uganda, produced more pant hoots on days when there were frequent changes in the male, but not female, composition of the focal's party. This factor accounted for the largest amount of variation in pant hoot production, and we found that males were more likely to repeat a call prior to rather than after fusion with other males, suggesting that the calls facilitate fusion. Pant hoots therefore seem to play a pivotal role in regulating grouping dynamics in chimpanzees. Our study also shows that pant hooting was positively correlated with the rank of the caller, the presence of parous females in estrus, and the consumption of highquality food, suggesting that pant hoots signal social status or social bonds when between-male competition is high. This study supports the view that pant hoots fulfill a complex social function.
The production of structured and repetitive sounds by striking objects is a behavior found not on... more The production of structured and repetitive sounds by striking objects is a behavior found not only in humans, but also in a variety of animal species, including chimpanzees (Pan troglodytes). In this study we examined individual and social factors that may influence the frequency with which individuals engage in drumming behavior when producing long distance pant hoot vocalizations, and analyzed the temporal structure of those drumming bouts. Male chimpanzees from Budongo Forest, Uganda, drummed significantly more frequently during travel than feeding or resting and older individuals were significantly more likely to produce drumming bouts than younger ones. In contrast, we found no evidence that the presence of estrus females, high ranking males and
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