Vocal Abilities in a Group of Nonhuman Primates
Résumé
Human language is too complex to have emerged in the absence of any evolutionary precursors, which suggests that primitive forms of pre-linguistic communication can be found in animals. Whether this was based on acoustic or gestural communication is an ongoing debate. An argument against a vocal origin is the absence of vocal flexibility and complexity in non-human primates. However, human language is primarily a vocal behaviour, and vocal flexibility – as assessed by vocal plasticity, semanticity, compositionality, and intentional signalling - is not a uniquely human trait. Our research focuses on the precursors of the various types of vocal flexibility in forest guenons (Cercopithecus spp). The vocal tract of nonhuman primates is in principle capable of producing speech-like sounds (Riede et al 2005) and one puzzle is why nonhuman primates do not make greater use of this feature. Instead, primates produce a finite range of calls that develop under strong genetic control. Within some call types, however, some flexibility can be seen at the level of call morphology, as for example demonstrated by socially-determined vocal plasticity and vocal sharing in Campbell's monkey contact calls used in conversation-like socially controlled vocal exchanges (Lemasson & Hausberger 2004; Lemasson et al 2010). Second, many primates produce acoustically distinct calls to specific external events, including Diana and Campbell's monkeys. In both species, the adult males and females produce acoustically different alarm calls to the same predator (Ouattara et al 2009a), but calls are meaningful to others, both within and between species. Alarm calls are not only predator-specific but also vary depending on the modality by which the predator is discovered, i.e. the visual or acoustic domain. In Campbell's monkeys females produced a complex alarm call repertoire, although differences were found between captive and wild individuals. Captive ones did not produce predator-specific calls but had a unique call to humans (Ouattara et al 2009a). For males, we found a repertoire of six call types, which could be classified into different morphs, according to the frequency contour and whether calls were trailed by an acoustically invariable suffix. Suffixed calls carried a broader meaning than unsuffixed ones (Ouattara et al 2009b). The six calls were concatenated into context-specific call sequences, following basic combinatorial principles (Ouattara et al 2009c). In sum, the vocal abilities in guenons go significantly beyond the currently assumed default case for nonhuman primates. Flexibility can be seen at all relevant levels, including limited control over call morphology, conversational rules, ability to produce context-specific calls, and some basic combinatorial properties. The data are at odds with a gestural origins of language theory. Gestural signals do not appear to play a key role in these species, while vocal flexibility is seen in all key components despite the fact that they have split from the human line about 30 million years ago. Field playback experiments will be needed to confirm whether receivers utilise these rich patterns to guide their behavioural decisions. But even in the absence of such evidence data suggest that a strong dichotomy between human language and nonhuman primate communication may no longer be tenable in the vocal domain. The visually dense forest habitat may have played a key role in the evolution of advanced vocalisation skills.