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  • Publication
    Accès libre
    Singing behavior of white-handed gibbons (Hylobates lar) of Khao Yai National Park, Thailand
    Ce manuscrit de thèse est axé sur les capacités vocales singulières des gibbons à mains blanches (Hylobates lar) du parc national de Khao Yai, en Thaïlande. Afin d’appréhender les mécanismes sous-jacents de leur système de communication, ce projet de recherche a été mené dans le parc national de Khao Yai en Thaïlande, au sein d’une population de gibbons dans leur environnement naturel, sur 13 groupes habitués à la présence humaine. Les gibbons à mains blanches, aussi appelés gibbon lar, font partie de la famille des Hylobatidés. Ils sont phylogénétiquement, les parents les plus éloignés de l’Homme au sein de la super-famille des Hominoïdae. L’étude de leur système complexe de communication s’avère donc, d’un très grand intérêt. Ils possèdent un répertoire de notes conséquent, produisent des chants de longue durée et ont la capacité de produire différents types de chants - résultant de la combinaison de plusieurs notes – ces attributs faisant d’eux un sujet d’étude unique pour aider à répondre à des problématiques scientifiques, telle que les origines du langage humain. Une caractéristique bien connue concernant les capacités vocales des gibbons, réside dans la production quotidienne et en synchronie, par le couple adulte mâle et femelle de chaque groupe, de chants stéréotypés longs et puissants ; qu’on appelle duos. On suppose que ces chants ont pour fonctions principales le renforcement du lien entre les individus du couple adulte, ainsi que la défense du territoire et du partenaire, cependant, ces hypothèses n’ont jamais été explorées de manière détaillée. Un autre aspect de leur production vocale concerne la production de chants prédateurs, qui, par le biais d’une précédente étude, présentent des différences structurelles notables qui les différencient des duos. Sur ce point également, nous en savons encore que très peu au-delà de ce postulat de base. L’objectif premier de ce projet de recherche était d’étudier les échanges interactifs de duos entre groupes de gibbons et d’explorer le rôle de la production de ces duos en tant que réseau de communication territorial complexe. Les duos sont généralement contagieux, se propageant aux groupes voisins, entrainant des vagues de duos dans la population. Cette étude a permis de démontrer que les duos émis en réponse à un duo précédant pouvaient être échangés en alternance de manière non-agressive, c’est-à-dire que les groupes vont temporiser le début de leur propre chant de sorte que le groupe précédent puisse terminer son duo sans être interrompu. Ces réponses pouvaient aussi être échangées de manière compétitive, en contre-chant, et dans ce cas les groupes vont initier leur propre duo par-dessus le duo précédent. Afin d’étudier ce phénomène, j’ai analysé les mécanismes régissant les échanges de duos entre deux groupes consécutifs. Les résultats obtenus ont montré que la distance entre les groupes, la composition sociale du groupe, et surtout le lien de parenté génétique entre les individus des deux groupes, avaient un effet significatif sur les modalités d’échange de duos entre groupes (contre-chant ou alternance). Cette étude a ainsi permis de mettre en évidence le rôle de la parenté dans les interactions vocales entre groupes, facilitant les échanges de manière non compétitive. La deuxième partie de ce projet de recherche est axée sur la production de chants prédateurs, et plus spécifiquement, sur la manière qu’ont les gibbons d’assembler différemment les mêmes éléments acoustiques de base de manière contextuelle, d’un type de chant à un autre. Je me suis donc intéressée à la façon dont les gibbons utilisent leurs chants sophistiqués pour encoder des événements externes et aux inférences que les individus réceptionnant ce signal peuvent extraire de ces vocalisations. De précédentes observations ont relaté la capacité des groupes de gibbons à répondre par le type de chant correspondant, c’est-à-dire à produire un duo en réponse à un duo émis par un groupe voisins, et de la même façon, à répondre par un chant prédateur en réponse à un autre chant prédateur. Ce phénomène se produisant en l’absence de contact visuel, sous-tend que les gibbons peuvent décoder et donc déduire la signification des différents types de chants produits par d’autres groupes, similaire à la façon dont les humains utilisent le langage pour communiquer et se comprendre. Afin d’étudier leurs capacités à produire des chants spécifiques et leurs capacités d’inférence, j’ai mené des expériences de playback et de présentation de modèles de prédateurs. Dans la première étude, j’ai diffusé, via un haut-parleur, des chants prédateurs ou des duos à différents groupes, et collecté leurs réponses vocales et comportementales. Dans la deuxième étude, j’ai présenté à plusieurs groupes différents modèles prédateurs, la panthère nébuleuse et le python réticulé, et de la même façon, j’ai collecté leurs réponses vocales et comportementales. Pour ce qui est des expériences de playback, les résultats ont démontré que les gibbons pouvaient discriminer et extraire des informations différentes de deux types de chants, duos et chants prédateurs, en manifestant différentes réponses vocales et comportementales. Concernant les présentations de modèles prédateurs, une étude précédente a montré que le modèle panthère suscitait chez les gibbons, et de manière fiable, de fortes réponses vocales et des comportements anti-prédateurs, contrairement au modèle python. Cette fois, le modèle de python a aussi provoqué de fortes réponses vocales et des comportements anti-prédateurs. Mais surtout, les analyses acoustiques ont montré que les divers types de chants produits pouvaient être différenciés par plusieurs structures acoustiques et arrangements de séquences, en fonction du type du type de prédateur, un critère de base de production de signaux d’alarmes référentiels. Les Hylobatidés, comprenant les gibbons, sont les premiers à avoir divergé de la famille des Hominoïdae. Leur position phylogénétique et leurs capacité vocales complexes, font des gibbons à mains blanches une espèce particulièrement prometteuse pour tenter de comprendre et d’expliquer comment les primates non-humains ont la capacité de coder leurs productions vocales en fonction du contexte, et les inférences qu’ils peuvent extraire de ces vocalisations. Ce projet a pour but d’apporter des données et des éléments pertinents, afin de mieux comprendre les conditions qui ont pu mener à l’émergence et à la complexité du langage humain. ABSTRACT This thesis manuscript focusses on the remarkable vocal abilities of white-handed gibbons (Hylobates lar) of Khao Yai National Park, Thailand. To understand the mechanisms underlying the acoustic communication system of these gibbons, I conducted my research project in Khao Yai National Park in Thailand, on 13 wild groups. White-handed gibbons, also known as lar gibbons, part of the Hylobatidae family, are the evolutionarily most distant relatives of humans within the hominoid lineage and are of interest regarding their complex vocal communication system. Their large vocal repertoire, lengthy songs and abilities to produce different song types - resultant from multiple note assemblages - make them an exceptional study system to address scientific questions, also about the origin of human vocal communication. A first well-known feature is that the mated pair sings stereotyped, loud and long songs on a daily basis and in a synchronised fashion; the duet songs. These songs are thought to function primarily in pair-bonding and mate and territorial defence, but these hypotheses have never been investigated in much depth. Another well-known feature is that gibbon groups also sing in response to predators and from previous research it became apparent that predator songs are assembled in different ways compared to duet songs. Here also, not much has been done beyond these basic findings. A first goal of this research was therefore to investigate the intergroup singing interactions and explore the role of duet songs as a complex territorial communication network. Duet songs often spread to neighbouring groups, resulting in song waves. In my research I have been able to show that duet responses can either non-aggressive turn-taking exchanges, where groups delay their song onset so that the preceding gibbon group can end their duet without interruption; alternatively, duet responses can be competitive, counter-singing exchanges, in which case the song of the preceding group is temporally overlapped by the subsequent duet song. To investigate this phenomenon, I examined the mechanisms underlying the duet song production of two subsequent groups. Results showed that intergroup distance, group composition and, most importantly, genetic relatedness had a significant impact on whether gibbons opted for competitive or non-aggressive duet exchanges with their neighbours. The study thus shows that kinship plays a role in non-aggressive interactions between groups. The second part of my research focusses on gibbon songs produced in response to predators, particularly how the same basic song units are assembled in context-specific ways. I was interested in how gibbons use their elaborate songs to encode external events and what inferences listeners can make from these vocalizations. Previous observations have shown that neighbouring groups respond to songs with the matching type, i.e. neighbours’ duets by their own duets, but also predator songs by their own predator songs. This happens in the absence of visual contact, suggesting that gibbons can decode and understand the meaning of each other’s songs, similar to how humans use language to address and understand each other. To investigate these coding and inference abilities in gibbons, I conducted playback and predator model experiments. In a first study, I broadcasted predator and duet songs to different groups and recorded vocal and behavioural responses. In a second study, I presented two predator types, the clouded leopard and the reticulated python, and again recorded subjects’ vocal and behavioural responses. Regarding the playback experiments, I found that gibbons could discriminate and infer the meaning of different song types (i.e. duet song and predator song) by displaying different vocal and behavioural responses. Regarding predator model presentations, a previous study showed that the leopard model reliably elicited strong vocal and anti-predatory behavioural responses, unlike the python model. This time, the life-sized python model also elicited strong singing and anti-predator responses, in contrast to previous research. More importantly, gibbons arranged song elements differently and songs could be distinguished regarding several acoustic components and sequential arrangements according to the type of predator encountered, a basic requirement for referential alarm calling behaviour. The basal position of gibbons in the Hominoïdae family combined with their usually complex vocal abilities make white-handed gibbons a particularly promising species to address questions of how nonhuman primates code external events in their vocal signals and what inferences they can make from these vocalizations. This kind of data will be relevant to better understand the conditions that might have led to the emergence and current complexity of human vocal communication, as seen in spoken language.
  • Publication
    Métadonnées seulement
    Predation affects alarm call usage in female Diana monkeys (Cercopithecus diana diana)
    Diana monkeys produce acoustically distinct calls to a number of external events, including different types of predators. In a previous study, we found population-wide differences in male alarm call production in Taï Forest, Ivory Coast, and on Tiwai Island, Sierra Leone, mostly likely originating from differences in predator experience. In Taï Forest, leopards (Panthera pardus) are common but on Tiwai Island they have been absent for decades, while the predation pressure from crowned eagles (Stephanoaetus coronatus) has been similar. To further evaluate the impact of predator experience, we here analyse the vocal behaviour of female Diana monkeys in both habitats. Female Diana monkeys produce predator-specific alarm calls, alert calls and contact calls in response to predators, suggesting that their calls serve in a broader range of functions compared to males. Results showed that females produced the same call types at both sites, despite the differences in predator fauna. Regarding call usage, leopard alarm calls were extremely rare on Tiwai Island, but not in Taï Forest, whereas we found no differences in eagle alarm call production. When comparing response latencies, Tiwai females were slower to respond to both predators compared to Taï females. Finally, we found no habitat-specific acoustic differences in the alert and predator-specific alarm calls, but significant differences in frequency-based parameters of contact calls. Overall, our results suggest that ontogenetic experience can affect primate vocal behaviour in both usage and acoustic structure but that the way in which particular call types are affected may be closely linked to function.
  • Publication
    Accès libre
    A forest monkey’s alarm call series to predator models
    (2008)
    Arnold, Kate
    ;
    Pohlner, Yvonne
    ;
    Some non-human primates produce acoustically distinct alarm calls to different predators, such as eagles or leopards. Recipients respond to these calls as if they have seen the actual predator, which has led to the notion of functionally referential alarm calls. However, in a previous study with free-ranging putty-nosed monkeys (Cercopithecus nictitans martini), we demonstrated that callers produced two acoustically distinct alarm calls to eagle shrieks and leopard growls, but both alarm calls were given to both predators. We can think of two basic explanations for this surprising result, a methodological and theoretical one. Firstly, acoustic predator models may not always be suitable to test alarm call behaviour in primates, sometimes causing uncharacteristic behaviour. Secondly, referential alarm calling may not be a universal feature of primate alarm call systems. Considering the methodological and theoretical importance of these possibilities, we conducted a follow-up study using life-sized leopard, eagle, and human models on the same population and compared the resulting vocal responses to those given to acoustic predator models. We compared the alarm call series given to each of these predator model types and found a considerable degree of consistency suggesting that the mode of presentation did not affect anti-predator calling strategies. However, evidence for audience effects on calling behaviour was inconclusive. While it appears that predator class is reliably encoded by different call series types irrespective of the mode of presentation, observations of these same call series given in non-predatory contexts indicate that predator class is unlikely to be the relevant organising principle underlying the alarm-calling behaviour in this species. We conclude by offering an alternative, non-referential, account of the alarmcalling system exhibited by this species.
  • Publication
    Accès libre
    Predator-specific alarm calls in Campbell’s monkeys,Cercopithecus campbelli
    One of the most prominent behavioural features of many forest primates are the loud calls given by the adult males. Early observational studies repeatedly postulated that these calls function in intragroup spacing or intergroup avoidance. More recent field experiments with Diana monkeys (Cercopithecus diana) of Ta Forest, Ivory Coast, have clearly shown that loud male calls function as predator alarm calls because calls reliably (1) label different predator classes and (2) convey semantic information about the predator type present. Here, I test the alarm call hypothesis another primate, the Campbell’s monkey (C. campbelli). Like Diana monkeys, male Campbell’s monkeys produce conspicuous loud calls to crowned hawk eagles (Stephanoaetus coronatus) and leopards (Panthera pardus), two of their main predators. Playback experiments showed that monkeys responded to the predator category represented by the different playback stimuli, regardless of whether they consisted of (1) vocalisations of the actual predators (crowned hawk eagle shrieks or leopard growls), (2) alarm calls to crowned hawk eagles or leopards given by other male Campbell’s monkeys or (3) alarm calls to crowned hawk eagles or leopards given by sympatric male Diana monkeys. These experiments provide further evidence that non-human primates have evolved the cognitive capacity to produce and respond to referential labels for external events.
  • Publication
    Accès libre
    The alarm call system of wild black-fronted titi monkeys, Callicebus nigrifrons
    Cäsar, Cristiane
    ;
    Byrne, Richard W
    ;
    Young, Robert J
    ;
    Upon encountering predators, many animals produce specific vocalisations that alert others and sometimes dissuade the predators from hunting. Callicebus monkeys are known for their large vocal repertoire, but little is known about the function and meaning of most call types. We recorded a large number of natural predator responses from five different groups of black-fronted titi monkeys in their Atlantic forest habitat in South Eastern Brazil. When detecting predatory threats, adult group members responded with call sequences that initially consisted of two brief, high-pitched calls with distinct frequency contours. Call A was mainly given to raptors but also to predatory capuchin monkeys and other threats within the canopy, while call B was given to predatory or non-predatory disturbances on the ground. In later parts of the sequences, we also recorded a high-pitched unmodulated call C and various low-pitched loud calls. Results therefore suggest that calls A and B provide listeners with rapid and reliable information about the general classes of danger experienced by the caller, while obtaining more specific information through other call types and combinations and behavioural responses. We discuss these findings in relation to current evolutionary theory of primate communication.
  • Publication
    Accès libre
    Speed of call delivery is related to context and caller identity in Campbell’s monkey males
    Lemasson, Alban
    ;
    Ouattara, Karim
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    Bouchet, Hélène
    ;
    Call rate can be a salient feature in animal communication. Depending on the species, different psychological variables appear to influence call rates but the exact nature of these relationships remains poorly explored. Here, we demonstrate for free-ranging Campbell's monkeys that the call rates of four different alarm series (termed H, K, K+, and B series) vary systematically as a function of context, associated behaviour, and identity of the caller. K+ series were given more rapidly to predation than non- predation events, K+ and K series more rapidly to visual than auditory predator detection, and H series more rapidly while counterattacking an eagle than staying put. Finally, there were individual differences in B series, suggesting that call rate potentially provides listeners with cues about the caller's anti-predator behaviour, event type experienced, and his identity.