Isochronous singing in 3 crested gibbon species (Nomascus spp.).

The search for common characteristics between the musical abilities of humans and other animal species is still taking its first steps. One of the most promising aspects from a comparative point of view is the analysis of rhythmic components, which are crucial features of human communicative performance but also well-identifiable patterns in the vocal displays of other species. Therefore, the study of rhythm is becoming essential to understand the mechanisms of singing behavior and the evolution of human communication. Recent findings provided evidence that particular rhythmic structures occur in human music and some singing animal species, such as birds and rock hyraxes, but only 2 species of nonhuman primates have been investigated so far (Indri indri and Hylobates lar). Therefore, our study aims to consistently broaden the list of species studied regarding the presence of rhythmic categories. We investigated the temporal organization in the singing of 3 species of crested gibbons (Nomascus gabriellae, Nomascus leucogenys, and Nomascus siki) and found that the most prominent rhythmic category was isochrony. Moreover, we found slight variation in songs' tempo among species, with N. gabriellae and N. siki singing with a temporal pattern involving a gradually increasing tempo (a musical accelerando), and N. leucogenys with a more regular pattern. Here, we show how the prominence of a peak at the isochrony establishes itself as a shared characteristic in the small apes considered so far.

Isochrony as ancestral condition to call and song in a primate.

Animal songs differ from calls in function and structure, and have comparative and translational value, showing similarities to human music. Rhythm in music is often distributed in quantized classes of intervals known as rhythmic categories. These classes have been found in the songs of a few nonhuman species but never in their calls. Are rhythmic categories song-specific, as in human music, or can they transcend the song-call boundary? We analyze the vocal displays of one of the few mammals producing both songs and call sequences: Indri indri. We test whether rhythmic categories (a) are conserved across songs produced in different contexts, (b) exist in call sequences, and (c) differ between songs and call sequences. We show that rhythmic categories occur across vocal displays. Vocalization type and function modulate deployment of categories. We find isochrony (1:1 ratio, like the rhythm of a ticking clock) in all song types, but only advertisement songs show three rhythmic categories (1:1, 1:2, 2:1 ratios). Like songs, some call types are also isochronous. Isochrony is the backbone of most indri vocalizations, unlike human speech, where it is rare. In indri, isochrony underlies both songs and hierarchy-less call sequences and might be ancestral to both.

Erratum to: Born to sing! Song development in a singing primate.

[This corrects the article DOI: 10.1093/cz/zoab018.].

Who you live with and what you duet for: a review of the function of primate duets in relation to their social organization.

Duets are one of the most fascinating displays in animal vocal communication, where two animals fine-tune the timing of their emissions to create a coordinated signal. Duetting behavior is widespread in the animal kingdom and is present in insects, birds, and mammals. Duets are essential to regulate activities within and between social units. Few studies assessed the functions of these vocal emissions experimentally, and for many species, there is still no consensus on what duets are used for. Here, we reviewed the literature on the function of duets in non-human primates, investigating a possible link between the social organization of the species and the function of its duetting behavior. In primates and birds, social conditions characterized by higher promiscuity might relate to the emergence of duetting behavior. We considered both quantitative and qualitative studies, which led us to hypothesize that the shift in the social organization from pair living to a mixed social organization might have led to the emergence of mate defense and mate guarding as critical functions of duetting behavior. Territory/resource ownership and defense functions are more critical in obligate pair-living species. Finally, we encourage future experimental research on this topic to allow the formulation of empirically testable predictions.

"The song remains the same": not really! Vocal flexibility in the song of the indris.

In studying communicative signals, we can think of flexibility as a necessary correlate of creativity. Flexibility enables animals to find practical solutions and appropriate behaviors in mutable situations. In this study, we aimed to quantify the degree of flexibility in the songs of indris (Indri indri), the only singing lemur, using three different metrics: Jaro Distance, normalized diversity, and entropy. We hypothesized that the degree and the co-variation of the flexibility of indris singing together would vary according to their status and sex. We found that dominant females were more flexible than dominant males when concatenating elements into strings (element concatenation). The number of different elements in a song contribution normalized by the contribution length (contribution diversity) of dominant individuals positively co-varied for seven duetting pairs. Non-dominant individuals were more variable in element concatenation than dominant individuals, and they were more diverse in phrase type than dominant females. Independently from sex and status, individual contributions did not differ in entropy (a measure of the predictability of contributions). These results corroborate previous findings regarding the dimorphism by sex and by status of individual contributions to songs. Thus, they shed light on the presence and expression of flexibility in the behavior of a non-human primate species. Indeed, they potentially show an effect of social features in shaping vocal flexibility, which underlies many communication systems, including human language. We speculate that this degree of flexibility may account for creativity.

Vocal tract shape variation contributes to individual vocal identity in African penguins.

Variation in formant frequencies has been shown to affect social interactions and sexual competition in a range of avian species. Yet, the anatomical bases of this variation are poorly understood. Here, we investigated the morphological correlates of formants production in the vocal apparatus of African penguins. We modelled the geometry of the supra-syringeal vocal tract of 20 specimens to generate a population of virtual vocal tracts with varying dimensions. We then estimated the acoustic response of these virtual vocal tracts and extracted the centre frequency of the first four predicted formants. We demonstrate that: (i) variation in length and cross-sectional area of vocal tracts strongly affects the formant pattern, (ii) the tracheal region determines most of this variation, and (iii) the skeletal size of penguins does not correlate with the trachea length and consequently has relatively little effect on formants. We conclude that in African penguins, while the variation in vocal tract geometry generates variation in resonant frequencies supporting the discrimination of conspecifics, such variation does not provide information on the emitter's body size. Overall, our findings advance our understanding of the role of formant frequencies in bird vocal communication.

Singing more, singing harsher: occurrence of nonlinear phenomena in a primate' song.

Nonlinear phenomena (NLP) in animal vocalizations arise from irregularities in the oscillation of the vocal folds. Various non-mutually exclusive hypotheses have been put forward to explain the occurrence of NLP, from adaptive to physiological ones. Non-human primates often display NLP in their vocalizations, yet the communicative role of these features, if any, is still unclear. We here investigate the occurrence of NLP in the song of a singing primate, the indri (Indri indri), testing for the effect of sex, age, season, and duration of the vocal display on their emission. Our results show that NLP occurrence in indri depends on phonation, i.e., the cumulative duration of all the units emitted by an individual, and that NLP have higher probability to be emitted in the later stages of the song, probably due to the fatigue indris may experience while singing. Furthermore, NLP happen earlier in the vocal display of adult females than in that of the adult males, and this is probably due to the fact that fatigue occurs earlier in the former because of a greater contribution within the song. Our findings suggest, therefore, that indris may be subjected to physiological constraints during the singing process which may impair the production of harmonic sounds. However, indris may still benefit from emitting NLP by strengthening the loudness of their signals for better advertising their presence to the neighboring conspecific groups.

There You Are! Automated Detection of Indris' Songs on Features Extracted from Passive Acoustic Recordings.

The growing concern for the ongoing biodiversity loss drives researchers towards practical and large-scale automated systems to monitor wild animal populations. Primates, with most species threatened by extinction, face substantial risks. We focused on the vocal activity of the indri (Indri indri) recorded in Maromizaha Forest (Madagascar) from 2019 to 2021 via passive acoustics, a method increasingly used for monitoring activities in different environments. We first used indris' songs, loud distinctive vocal sequences, to detect the species' presence. We processed the raw data (66,443 10-min recordings) and extracted acoustic features based on the third-octave band system. We then analysed the features extracted from three datasets, divided according to sampling year, site, and recorder type, with a convolutional neural network that was able to generalise to recording sites and previously unsampled periods via data augmentation and transfer learning. For the three datasets, our network detected the song presence with high accuracy (>90%) and recall (>80%) values. Once provided the model with the time and day of recording, the high-performance values ensured that the classification process could accurately depict both daily and annual habits of indris' singing pattern, critical information to optimise field data collection. Overall, using this easy-to-implement species-specific detection workflow as a preprocessing method allows researchers to reduce the time dedicated to manual classification.

Born to sing! Song development in a singing primate.

In animal vocal communication, the development of adult-like vocalization is fundamental to interact appropriately with conspecifics. However, the factors that guide ontogenetic changes in the acoustic features remain poorly understood. In contrast with a historical view of nonhuman primate vocal production as substantially innate, recent research suggests that inheritance and physiological modification can only explain some of the developmental changes in call structure during growth. A particular case of acoustic communication is the indris' singing behavior, a peculiar case among Strepsirrhine primates. Thanks to a decade of intense data collection, this work provides the first long-term quantitative analysis on song development in a singing primate. To understand the ontogeny of such a complex vocal output, we investigated juvenile and sub-adult indris' vocal behavior, and we found that young individuals started participating in the chorus years earlier than previously reported. Our results indicated that spectro-temporal song parameters underwent essential changes during growth. In particular, the age and sex of the emitter influenced the indris' vocal activity. We found that frequency parameters showed consistent changes across the sexes, but the temporal features showed different developmental trajectories for males and females. Given the low level of morphological sexual dimorphism and the marked differences in vocal behavior, we hypothesize that factors like social influences and auditory feedback may affect songs' features, resulting in high vocal flexibility in juvenile indris. This trait may be pivotal in a species that engages in choruses with rapid vocal turn-taking.