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.

Predation on gibbons is rarely observed in the wild. However, the gibbons' moderate body size and relatively small social groups suggest high vulnerability to predation. To assess the role of predation and to study their anti-predator behaviour, we presented visual predator models to nine groups of wild white-handed gibbons at Khao Yai National Park, Thailand. We measured subjects' immediate and delayed responses to four potential predators: tiger, clouded leopard, crested serpent eagle and reticulated python. Subjects reliably approached all four predators. In response to tigers and leopards, they additionally produced predator-specific songs and defecated copiously. In terms of delayed responses, distance between mated adults decreased, but only after exposure to the tiger model. In response to eagles and pythons, gibbons consistently vocalised, but this did not always include predator singing, and we found no long-term effects in overall activity or strata use. However, during 6 of 26 predator encounters, the gibbons produced songs with a structure that was intermediate between a duet song and a predator song more than 20 min after the predator encounter, indicating a long-term effect on their vocal behaviour. This study demonstrates that gibbons discriminate between different potential predators and respond to them with adaptive anti-predator behaviour, which include predator-specific vocal responses. We conclude that gibbons are not immune to predation and that terrestrial predators elicit consistent immediate and delayed anti-predation responses.

Using artificially synthesized stimuli, previous research has shown that cotton-top tamarin monkeys easily learn simple AB grammar sequences, but not the more complex AnBn sequences that require hierarchical structure. Humans have no trouble learning AnBn combinations. A more recent study, using similar artificially created stimuli, showed that there is a neuroanatomical difference in the brain between these two kinds of arrays. While the simpler AB sequences recruit the frontal operculum, the AnBn array recruits the phylogenetically newer Broca’s area. We propose that on close inspection, reported vocal repertoires of Old World Monkeys show that these nonhuman primates are capable of calls that have two items in them, but never more than two. These are simple AB sequences, as predicted by previous research. In addition, we suggest the two-item call cannot be the result of a combinatorial operation that we see in human language, where the recursive operation of Merge allows for a potentially infinite array of structures. In our view, the two-item calls of nonhuman primates result from a dual-compartment frame into which each of the calls can fit without having to be combined by an operation such as Merge.

BackgroundClose range calls are produced by many animals during intra-specific interactions, such as during home range defence, playing, begging for food, and directing others. In this study, we investigated the most common close range vocalisation of lar gibbons (Hylobates lar), the ‘hoo’ call. Gibbons and siamangs (family Hylobatidae) are known for their conspicuous and elaborate songs, while quieter, close range vocalisations have received almost no empirical attention, perhaps due to the difficult observation conditions in their natural forest habitats.ResultsWe found that ‘hoo’ calls were emitted by both sexes in a variety of contexts, including feeding, separation from group members, encountering predators, interacting with neighbours, or as part of duet songs by the mated pair. Acoustic analyses revealed that ‘hoo’ calls varied in a number of spectral parameters as a function of the different contexts. Males’ and females’ ‘hoo’ calls showed similar variation in these context-specific parameter differences, although there were also consistent sex differences in frequency across contexts.ConclusionsOur study provides evidence that lar gibbons are able to generate significant, context-dependent acoustic variation within their main social call, which potentially allows recipients to make inferences about the external events experienced by the caller. Communicating about different events by producing subtle acoustic variation within some call types appears to be a general feature of primate communication, which can increase the expressive power of vocal signals within the constraints of limited vocal tract flexibility that is typical for all non-human primates. In this sense, this study is of direct relevance for the on-going debate about the nature and origins of vocally-based referential communication and the evolution of human speech.