Chimpanzee vowel-like sounds and voice quality suggest formant space expansion through the hominoid lineage.

The origins of human speech are obscure; it is still unclear what aspects are unique to our species or shared with our evolutionary cousins, in part due to a lack of a common framework for comparison. We asked what chimpanzee and human vocal production acoustics have in common. We examined visible supra-laryngeal articulators of four major chimpanzee vocalizations (hoos, grunts, barks, screams) and their associated acoustic structures, using techniques from human phonetic and animal communication analysis. Data were collected from wild adult chimpanzees, Taï National Park, Ivory Coast. Both discriminant and principal component classification procedures revealed classification of call types. Discriminating acoustic features include voice quality and formant structure, mirroring phonetic features in human speech. Chimpanzee lip and jaw articulation variables also offered similar discrimination of call types. Formant maps distinguished call types with different vowel-like sounds. Comparing our results with published primate data, humans show less F1-F2 correlation and further expansion of the vowel space, particularly for [i] sounds. Unlike recent studies suggesting monkeys achieve human vowel space, we conclude from our results that supra-laryngeal articulatory capacities show moderate evolutionary change, with vowel space expansion continuing through hominoid evolution. Studies on more primate species will be required to substantiate this. This article is part of the theme issue 'Voice modulation: from origin and mechanism to social impact (Part II)'.

Rethinking the frequency code: a meta-analytic review of the role of acoustic body size in communicative phenomena.

The widely cited frequency code hypothesis attempts to explain a diverse range of communicative phenomena through the acoustic projection of body size. The set of phenomena includes size sound symbolism (using /i/ to signal smallness in words such as teeny), intonational phonology (using rising contours to signal questions) and the indexing of social relations via vocal modulation, such as lowering one's voice pitch to signal dominance. Among other things, the frequency code is commonly interpreted to suggest that polite speech should be universally signalled via high pitch owing to the association of high pitch with small size and submissiveness. We present a cross-cultural meta-analysis of polite speech of 101 speakers from seven different languages. While we find evidence for cross-cultural variation, voice pitch is on average lower when speakers speak politely, contrary to what the frequency code predicts. We interpret our findings in the light of the fact that pitch has a multiplicity of possible communicative meanings. Cultural and contextual variation determines which specific meanings become manifest in a specific interactional context. We use the evidence from our meta-analysis to propose an updated view of the frequency code hypothesis that is based on the existence of many-to-many mappings between speech acoustics and communicative interpretations. This article is part of the theme issue 'Voice modulation: from origin and mechanism to social impact (Part I)'.

Chest beats as an honest signal of body size in male mountain gorillas (Gorilla beringei beringei).

Acoustic signals that reliably indicate body size, which usually determines competitive ability, are of particular interest for understanding how animals assess rivals and choose mates. Whereas body size tends to be negatively associated with formant dispersion in animal vocalizations, non-vocal signals have received little attention. Among the most emblematic sounds in the animal kingdom is the chest beat of gorillas, a non-vocal signal that is thought to be important in intra and inter-sexual competition, yet it is unclear whether it reliably indicates body size. We examined the relationship among body size (back breadth), peak frequency, and three temporal characteristics of the chest beat: duration, number of beats and beat rate from sound recordings of wild adult male mountain gorillas. Using linear mixed models, we found that larger males had significantly lower peak frequencies than smaller ones, but we found no consistent relationship between body size and the temporal characteristics measured. Taken together with earlier findings of positive correlations among male body size, dominance rank and reproductive success, we conclude that the gorilla chest beat is an honest signal of competitive ability. These results emphasize the potential of non-vocal signals to convey important information in mammal communication.

Response to Garcia and Dunn.

Garcia and Dunn [1] raise some interesting and valuable points regarding our recent paper in Current Biology[2]. As Garcia and Dunn [1] point out, cross-species variation in vocal and anatomical relations allows for the identification of relevant outliers from the body size - fundamental frequency (f0) regression. However, this depends on the premise that the chosen or available f0 and body size values are typical of the species. A motivation for our study [2] was in part to improve the accuracy of such estimates by providing more data per species compared to previous studies. We address each point of their critique by controlling for cross-species body size variation using body weights for chimpanzees (Pan troglodytes) and bonobos (Pan paniscus), addressing potential call variation in different subspecies of Pan troglodytes, measuring minimum f0 as well as maximum f0 and possible effects caused by different larynx fixation methods.

Reducing language to rhythm: Amazonian Bora drummed language exploits speech rhythm for long-distance communication.

Many drum communication systems around the world transmit information by emulating tonal and rhythmic patterns of spoken languages in sequences of drumbeats. Their rhythmic characteristics, in particular, have not been systematically studied so far, although understanding them represents a rare occasion for providing an original insight into the basic units of speech rhythm as selected by natural speech practices directly based on beats. Here, we analyse a corpus of Bora drum communication from the northwest Amazon, which is nowadays endangered with extinction. We show that four rhythmic units are encoded in the length of pauses between beats. We argue that these units correspond to vowel-to-vowel intervals with different numbers of consonants and vowel lengths. By contrast, aligning beats with syllables, mora or only vowel length yields inconsistent results. Moreover, we also show that Bora drummed messages conventionally select rhythmically distinct markers to further distinguish words. The two phonological tones represented in drummed speech encode only few lexical contrasts. Rhythm thus appears to crucially contribute to the intelligibility of drummed Bora. Our study provides novel evidence for the role of rhythmic structures composed of vowel-to-vowel intervals in the complex puzzle concerning the redundancy and distinctiveness of acoustic features embedded in speech.

Higher fundamental frequency in bonobos is explained by larynx morphology.

Acoustic signals, shaped by natural and sexual selection, reveal ecological and social selection pressures [1]. Examining acoustic signals together with morphology can be particularly revealing. But this approach has rarely been applied to primates, where clues to the evolutionary trajectory of human communication may be found. Across vertebrate species, there is a close relationship between body size and acoustic parameters, such as formant dispersion and fundamental frequency (f0). Deviations from this acoustic allometry usually produce calls with a lower f0 than expected for a given body size, often due to morphological adaptations in the larynx or vocal tract [2]. An unusual example of an obvious mismatch between fundamental frequency and body size is found in the two closest living relatives of humans, bonobos (Pan paniscus) and chimpanzees (Pan troglodytes). Although these two ape species overlap in body size [3], bonobo calls have a strikingly higher f0 than corresponding calls from chimpanzees [4]. Here, we compare acoustic structures of calls from bonobos and chimpanzees in relation to their larynx morphology. We found that shorter vocal fold length in bonobos compared to chimpanzees accounted for species differences in f0, showing a rare case of positive selection for signal diminution in both bonobo sexes.