Social factors drive vocal exchanges in red-capped mangabeys (Cercocebus torquatus).

Of all the calls made by non-human primates, the function of short-distance contact calls has largely remained to be determined. These calls are the most frequent in the repertoire and are most often exchanged between individuals in a non-random way. To our knowledge, no study has ever examined how vocal exchanges are structured in red-capped mangabeys (Cercocebus torquatus), a semi-terrestrial monkey living in the African forest with a complex semi-tolerant/semi-despotic social system. Our goal was to assess the organization of contact call exchanges in this species and their relationship with individual and social factors such as age, affinity, and hierarchy. Therefore, we observed several captive groups of red-capped mangabeys and collected data on vocal behavior, as well as grooming, agonistic behavior, and spatial proximity. We defined a vocal exchange in this species as a series of contact calls made by two or more individuals within a maximum inter-caller interval of 2 s. At the individual level, the higher the individual's hierarchical rank, the less they initiated exchanges. Furthermore, the most socially integrated individuals had a longer average response time than the less integrated ones. At the dyadic level, preferred exchange partners were individuals often observed near one other or individuals most distant in age. Also, the further apart two individuals were in the dominance hierarchy, the shorter the response time. Our results support both the social bonding hypothesis and a modulating key role of the dominance hierarchy on the social use of contact calls, which is in line with the social style of this species.

Suffixation influences receivers' behaviour in non-human primates.

Compared to humans, non-human primates have very little control over their vocal production. Nonetheless, some primates produce various call combinations, which may partially offset their lack of acoustic flexibility. A relevant example is male Campbell's monkeys (Cercopithecus campbelli), which give one call type ('Krak') to leopards, while the suffixed version of the same call stem ('Krak-oo') is given to unspecific danger. To test whether recipients attend to this suffixation pattern, we carried out a playback experiment in which we broadcast naturally and artificially modified suffixed and unsuffixed 'Krak' calls of male Campbell's monkeys to 42 wild groups of Diana monkeys (Cercopithecus diana diana). The two species form mixed-species groups and respond to each other's vocalizations. We analysed the vocal response of male and female Diana monkeys and overall found significantly stronger vocal responses to unsuffixed (leopard) than suffixed (unspecific danger) calls. Although the acoustic structure of the 'Krak' stem of the calls has some additional effects, subject responses were mainly determined by the presence or the absence of the suffix. This study indicates that suffixation is an evolved function in primate communication in contexts where adaptive responses are particularly important.

Dance displays in gibbons: biological and linguistic perspectives on structured, intentional, and rhythmic body movement.

Female crested gibbons (genus Nomascus) perform conspicuous sequences of twitching movements involving the rump and extremities. However, these dances have attracted little scientific attention and their structure and meaning remain largely obscure. Here we analyse close-range video recordings of captive crested gibbons, extracting descriptions of dance in four species (N. annamensis, N. gabriellae, N. leucogenys and N. siki). In addition, we report results from a survey amongst relevant professionals clarifying behavioural contexts of dance in captive and wild crested gibbons. Our results demonstrate that dances in Nomascus represent a common and intentional form of visual communication restricted to sexually mature females. Whilst primarily used as a proceptive signal to solicit copulation, dances occur in a wide range of contexts related to arousal and/or frustration in captivity. A linguistically informed view of this sequential behaviour demonstrates that movement within dances is organized in groups and follows an isochronous rhythm - patterns not described for visual displays in other non-human primates. We argue that applying the concept of dance to gibbons allows us to expand our understanding of communication in non-human primates and to develop hypotheses on the rules and regularities characterising it. We propose that crested gibbon dances likely evolved from less elaborate rhythmic proceptive signals, similar to those found in siamangs. Although dance displays in humans and crested gibbons share a number of key characteristics, they cannot be assumed to be homologous. Nevertheless, gibbon dances represent a striking model behaviour to investigate the use of complex gestural signals in hominoid primates.

Minimal Compositionality versus Bird Implicatures: two theories of ABC-D sequences in Japanese tits.

It was argued in a series of experimental studies that Japanese tits (Parus minor) have an ABC call that has an alert function, a D call that has a recruitment function, and an ABC-D call that is compositionally derived from ABC and D, and has a mobbing function. A key conclusion was that ABC-D differs from the combination of separate utterances of ABC and of D (e.g. as played by distinct but close loudspeakers). While the logic of the argument is arguably sound, no explicit rule has been proposed to derive the meaning of ABC-D from that of its parts. We compare two analyses. One posits a limited instance of semantic compositionality ('Minimal Compositionality'); the other does without compositionality, but uses instead a more sophisticated pragmatics ('Bird Implicatures'). Minimal Compositionality takes the composition of ABC and D to deviate only minimally from what would be found with two independent utterances: ABC means that 'there is something that licenses an alert', D means that 'there is something that licenses recruitment', and ABC-D means that 'there is something that licenses both an alert and recruitment'. By contrast, ABC and D as independent utterances yield something weaker, namely: 'there is something that licenses an alert, and there is something that licenses recruitment', without any 'binding' across the two utterances. The second theory, Bird Implicatures, only requires that ABC-D should be more informative than ABC, and/or than D. It builds on the idea, proposed for several monkey species, that a less-informative call competes with a more informative one (the 'Informativity Principle'): when produced alone, ABC and D trigger an inference that ABC-D is false. We explain how both Minimal Compositionality and Bird Implicatures could have evolved, and we compare the predictions of the two theories. Finally, we extend the discussion to some chimpanzee and meerkat sequences that might raise related theoretical problems.

Animal linguistics: a primer.

The evolution of language has been investigated by several research communities, including biologists and linguists, striving to highlight similar linguistic capacities across species. To date, however, no consensus exists on the linguistic capacities of non-human species. Major controversies remain on the use of linguistic terminology, analysis methods and behavioural data collection. The field of 'animal linguistics' has emerged to overcome these difficulties and attempt to reach uniform methods and terminology. This primer is a tutorial review of 'animal linguistics'. It describes the linguistic concepts of semantics, pragmatics and syntax, and proposes minimal criteria to be fulfilled to claim that a given species displays a particular linguistic capacity. Second, it reviews relevant methods successfully applied to the study of communication in animals and proposes a list of useful references to detect and overcome major pitfalls commonly observed in the collection of animal behaviour data. This primer represents a step towards mutual understanding and fruitful collaborations between linguists and biologists.

The ABC-D of animal linguistics: are syntax and compositionality for real?

In several animal species, an alarm call (e.g. ABC notes in the Japanese tit Parus minor) can be immediately followed by a recruitment call (e.g. D notes) to yield a complex call that triggers a third behaviour, namely mobbing. This has been taken to be an argument for animal syntax and compositionality (i.e. the property by which the meaning of a complex expression depends on the meaning of its parts and the way they are put together). Several additional discoveries were made across species. First, in some cases, animals respond with mobbing to the order alarm-recruitment but not to the order recruitment-alarm. Second, animals sometimes respond similarly to functionally analogous heterospecific calls they have never heard before, and/or to artificial hybrid sequences made of conspecific and heterospecific calls in the same order, thus adding an argument for the productivity of the relevant rules. We consider the details of these arguments for animal syntax and compositionality and argue that, with one important exception (Japanese tit ABC-D sequences), they currently remain ambiguous: there are reasonable alternatives on which each call is a separate utterance and is interpreted as such ('trivial compositionality'). More generally, we propose that future studies should argue for animal syntax and compositionality by explicitly pitting the target theory against two deflationary analyses: the 'only one expression' hypothesis posits that there is no combination in the first place, for example just a simplex ABCD call; while the 'separate utterances' hypothesis posits that there are separate expressions (e.g. ABC and D), but that they form separate utterances and are neither syntactically nor semantically combined.

Dominance style is a key predictor of vocal use and evolution across nonhuman primates.

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 vocalizations than their tolerant counterparts. Findings suggest primate signals are used and evolve in tandem with the nature of interactions that characterize individuals' social relationships.