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One of the most apparent discontinuities between non-human primate (primate) call communication and human speech concerns repertoire size. The former is essentially fixed to a limited number of innate calls, while the latter essentially consists of numerous learned components. Consequently, primates are thought to lack laryngeal control required to produce learned voiced calls. However, whether they may produce learned voiceless calls awaits investigation. Here, a case of voiceless call learning in primates is investigated--orangutan (Pongo spp.) whistling. In this study, all known whistling orangutans are inventoried, whistling-matching tests (previously conducted with one individual) are replicated with another individual using original test paradigms, and articulatory and acoustic whistle characteristics are compared between three orangutans. Results show that whistling has been reported for ten captive orangutans. The test orangutan correctly matched human whistles with significantly high levels of performance. Whistle variation between individuals indicated voluntary control over the upper lip, lower lip, and respiratory musculature, allowing individuals to produce learned voiceless calls. Results are consistent with inter- and intra-specific social transmission in whistling orangutans. Voiceless call learning in orangutans implies that some important components of human speech learning and control were in place before the homininae-ponginae evolutionary split.

The capacity of nonhuman primates to actively modify the acoustic structure of existing sounds or vocalizations in their repertoire appears limited. Several studies have reported population or community differences in the acoustical structure of nonhuman primate long distance calls and have suggested vocal learning as a mechanism for explaining such variation. In addition, recent studies on great apes have indicated that there are repertoire differences between populations. Some populations have sounds in their repertoire that others have not. These differences have also been suggested to be the result of vocal learning. On yet another level great apes can, after extensive human training, also learn some species atypical vocalizations. Here we show a new aspect of great ape vocal learning by providing data that an orangutan has spontaneously (without any training) acquired a human whistle and can modulate the duration and number of whistles to copy a human model. This might indicate that the learning capacities of great apes in the auditory domain might be more flexible than hitherto assumed.

The evolutionary origins of speech remain obscure. Recently, it was proposed that speech derived from monkey facial signals which exhibit a speech-like rhythm of ∼5 open-close lip cycles per second. In monkeys, these signals may also be vocalized, offering a plausible evolutionary stepping stone towards speech. Three essential predictions remain, however, to be tested to assess this hypothesis' validity; (i) Great apes, our closest relatives, should likewise produce 5Hz-rhythm signals, (ii) speech-like rhythm should involve calls articulatorily similar to consonants and vowels given that speech rhythm is the direct product of stringing together these two basic elements, and (iii) speech-like rhythm should be experience-based. Via cinematic analyses we demonstrate that an ex-entertainment orangutan produces two calls at a speech-like rhythm, coined “clicks” and “faux-speech.” Like voiceless consonants, clicks required no vocal fold action, but did involve independent manoeuvring over lips and tongue. In parallel to vowels, faux-speech showed harmonic and formant modulations, implying vocal fold and supralaryngeal action. This rhythm was several times faster than orangutan chewing rates, as observed in monkeys and humans. Critically, this rhythm was seven-fold faster, and contextually distinct, than any other known rhythmic calls described to date in the largest database of the orangutan repertoire ever assembled. The first two predictions advanced by this study are validated and, based on parsimony and exclusion of potential alternative explanations, initial support is given to the third prediction. Irrespectively of the putative origins of these calls and underlying mechanisms, our findings demonstrate irrevocably that great apes are not respiratorily, articulatorilly, or neurologically constrained for the production of consonant- and vowel-like calls at speech rhythm. Orangutan clicks and faux-speech confirm the importance of rhythmic speech antecedents within the primate lineage, and highlight potential articulatory homologies between great ape calls and human consonants and vowels. [ABSTRACT FROM AUTHOR]