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.

Evidence for compositionality in baboons (Papio papio) through the test case of negation.

Can non-human animals combine abstract representations much like humans do with language? In particular, can they entertain a compositional representation such as 'not blue'? Across two experiments, we demonstrate that baboons (Papio papio) show a capacity for compositionality. Experiment 1 showed that baboons can entertain negative, compositional, representations: they can learn to associate a cue with iconically related referents (e.g., a blue patch referring to all blue objects), but also to the complement set associated with it (e.g., a blue patch referring to all non-blue objects). Strikingly, Experiment 2 showed that baboons not only learn to associate a cue with iconically related referents, but can learn to associate complex cues (composed of the same cue and an additional visual element) with the complement object set. Thus, they can learn an operation, instantiated by this additional visual element, that can be compositionally combined with previously learned cues. These results significantly reduce any claim that would make the manipulation and combination of abstract representations a solely human privilege.

Impact of predator model presentation paradigms on titi monkey alarm sequences.

Abstract: Predator presentation experiments are widely used to investigate animal alarm vocalizations. They usually involve presentations of predator models or playbacks of predator calls, but it remains unclear whether the two paradigms provide similar results, a major limitation when investigating animal syntactic and semantic capacities. Here, we investigate whether visual and acoustic predator cues elicit different vocal reactions in black-fronted titi monkeys (Callicebus nigrifrons). We exposed six groups of wild titi monkeys to visual models or playbacks of vocalizations of raptor or felid. We characterized each group's vocal reactions using sequence parameters known to reliably encode predatory events in this species. We found that titi monkeys' vocal reactions varied with the predator species but also with the experimental paradigm: while vocal reactions to raptor vocalizations and models were similar, felid vocalizations elicited heterogeneous, different reactions from that given to felid models. We argue that subjects are not familiar with felid vocalizations, because of a lack of learning opportunities due to the silent behaviour of felids. We discuss the implication of these findings for the semantic capacities of titi monkeys. We finally recommend that playbacks of predator vocalizations should not be used in isolation but in combination with visual model presentations, to allow fine-grained analyses of the communication system of prey species. Significance statement: It is common to present prey species with predator models or predator calls to study their vocal reactions. The two paradigms are often used independently, but it remains unclear whether they provide similar results. Here, we studied the vocal reactions of titi monkeys to calls and models of raptors and felids. We show that titi monkeys seem to recognize the vocalizations of raptors but not those of felids. The study of the vocal reactions emitted when titi monkeys cannot clearly identify the threat allows us to draw accurate hypotheses about the meaning of titi monkeys' alarm utterances. We argue that playbacks of predator calls should be used in conjunction with model presentations, which can allow us to better investigate the information and the structure of the alarm systems. Supplementary Information: The online version contains supplementary material available at 10.1007/s00265-022-03250-1.

Dramatic decline in a titi monkey population after the 2016-2018 sylvatic yellow fever outbreak in Brazil.

Platyrrhini are highly vulnerable to the yellow fever (YF) virus. From 2016 to 2018, the Atlantic Forest of southeast Brazil faced its worst sylvatic YF outbreak in about a century, thought to have killed thousands of primates. It is essential to assess the impact of this epidemic on threatened primate assemblages to design effective conservation strategies. In this study, we assessed the impact of the 2016-2018 YF outbreak on a geographically isolated population of Near Threatened black-fronted titi monkeys (Callicebus nigrifrons) in two Atlantic Forest patches of the Santuário do Caraça, MG, Brazil. Extensive preoutbreak monitoring, conducted between 2008 and 2016, revealed that the home range and group sizes of the population remained stable. In 2016, the population size was estimated at 53-57 individuals in 11-12 groups. We conducted monitoring and playback surveys in 2019 and found that the population had decreased by 68% in one forest patch and completely vanished in the other, resulting in a combined decline of 80%. We discuss this severe loss of a previously stable population and conclude that it was highly likely caused by the YF outbreak. The remaining population is at risk of disappearing completely because of its small size and geographic isolation. A systematic population surveys of C. nigrifrons, along other sensible Platyrrhini species, is needed to re-evaluate their current conservation status.

Socially mediated overlap in vocal interactions between free-ranging black howler monkeys.

"Conversation rules" such as overlap avoidance and coordinated overlap have been reported in nonhuman animals, and seem to be adaptive responses to the requirements of social life. Some species display both patterns in an apparently flexible way, but the social factors mediating their respective usage remain poorly documented. We investigated the potential social factors guiding the usage of these temporal rules during collective howling in six free-ranging groups of black howler monkeys (Alouatta pigra). First, we found that adult males' vocalizations rarely overlap those of other callers while most adult females' calls overlap other members' calls, especially that of other females. Second, whereas some call types (notably affiliative calls) are typically emitted without overlap, roars (agonistic loud calls) overlap more frequently. Third, coordinated overlap is more frequent during intergroup competition. Our findings support the hypothesis that overlap avoidance and coordinated overlap are two different (here sex-related) vocal alliance social behaviors, at least for some nonhuman primates. More comparative investigations are now needed to explore further their evolutionary trajectories in this lineage.

Titi monkeys combine alarm calls to create probabilistic meaning.

Previous work suggested that titi monkeys Callicebus nigrifrons combine two alarm calls, the A- and B-calls, to communicate about predator type and location. To explore how listeners process these sequences, we recorded alarm call sequences of six free-ranging groups exposed to terrestrial and aerial predator models, placed on the ground or in the canopy, and used multimodel inference to assess the information encoded in the sequences. We then carried out playback experiments to identify the features used by listeners to react to the available information. Results indicated that information about predator type and location were encoded by the proportion of B-call pairs relative to all call pairs of the sequence (i.e., proportion of BB-grams). The results suggest that the meaning of the sequence is not conveyed in a categorical but probabilistic manner. We discuss the implications of these findings for current theories of animal communication and language evolution.