Sound Visualization Demonstrates Velopharyngeal Coupling and Complex Spectral Variability in Asian Elephants.

Sound production mechanisms set the parameter space available for transmitting biologically relevant information in vocal signals. Low-frequency rumbles play a crucial role in coordinating social interactions in elephants' complex fission-fusion societies. By emitting rumbles through either the oral or the three-times longer nasal vocal tract, African elephants alter their spectral shape significantly. In this study, we used an acoustic camera to visualize the sound emission of rumbles in Asian elephants, which have received far less research attention than African elephants. We recorded nine adult captive females and analyzed the spectral parameters of 203 calls, including vocal tract resonances (formants). We found that the majority of rumbles (64%) were nasally emitted, 21% orally, and 13% simultaneously through the mouth and trunk, demonstrating velopharyngeal coupling. Some of the rumbles were combined with orally emitted roars. The nasal rumbles concentrated most spectral energy in lower frequencies exhibiting two formants, whereas the oral and mixed rumbles contained higher formants, higher spectral energy concentrations and were louder. The roars were the loudest, highest and broadest in frequency. This study is the first to demonstrate velopharyngeal coupling in a non-human animal. Our findings provide a foundation for future research into the adaptive functions of the elephant acoustic variability for information coding, localizability or sound transmission, as well as vocal flexibility across species.

Acoustic structure and information content of trumpets in female Asian elephants (Elephas maximus).

Most studies on elephant vocal communication have focused on the low-frequency rumble, with less effort on other vocalization types such as the most characteristic elephant call, the trumpet. Yet, a better and more complete understanding of the elephant vocal system requires investigating other vocalization types and their functioning in more detail as well. We recorded adult female Asian elephants (Elephas maximus) at a private facility in Nepal and analyzed 206 trumpets from six individuals regarding their frequency, temporal and contour shape, and related acoustic parameters of the fundamental frequency. We also tested for information content regarding individuality and context. Finally, we recorded the occurrence of non-linear phenomena such as bifurcation, biphonation, subharmonics and deterministic chaos. We documented a mean fundamental frequency ± SD of 474 ± 70 Hz and a mean duration ± SD of 1.38 ± 1.46 s (Nindiv. = 6, Ncalls = 206). Our study reveals that the contour of the fundamental frequency of trumpets encodes information about individuality, but we found no evidence for trumpet subtypes in greeting versus disturbance contexts. Non-linear phenomena prevailed and varied in abundance among individuals, suggesting that irregularities in trumpets might enhance the potential for individual recognition. We propose that trumpets in adult female Asian elephants serve to convey an individual's identity as well as to signal arousal and excitement to conspecifics.

The multi-dimensional nature of vocal learning.

How learning affects vocalizations is a key question in the study of animal communication and human language. Parallel efforts in birds and humans have taught us much about how vocal learning works on a behavioural and neurobiological level. Subsequent efforts have revealed a variety of cases among mammals in which experience also has a major influence on vocal repertoires. Janik and Slater (Anim. Behav.60, 1-11. (doi:10.1006/anbe.2000.1410)) introduced the distinction between vocal usage and production learning, providing a general framework to categorize how different types of learning influence vocalizations. This idea was built on by Petkov and Jarvis (Front. Evol. Neurosci.4, 12. (doi:10.3389/fnevo.2012.00012)) to emphasize a more continuous distribution between limited and more complex vocal production learners. Yet, with more studies providing empirical data, the limits of the initial frameworks become apparent. We build on these frameworks to refine the categorization of vocal learning in light of advances made since their publication and widespread agreement that vocal learning is not a binary trait. We propose a novel classification system, based on the definitions by Janik and Slater, that deconstructs vocal learning into key dimensions to aid in understanding the mechanisms involved in this complex behaviour. We consider how vocalizations can change without learning, and a usage learning framework that considers context specificity and timing. We identify dimensions of vocal production learning, including the copying of auditory models (convergence/divergence on model sounds, accuracy of copying), the degree of change (type and breadth of learning) and timing (when learning takes place, the length of time it takes and how long it is retained). We consider grey areas of classification and current mechanistic understanding of these behaviours. Our framework identifies research needs and will help to inform neurobiological and evolutionary studies endeavouring to uncover the multi-dimensional nature of vocal learning. This article is part of the theme issue 'Vocal learning in animals and humans'.

A novel theory of Asian elephant high-frequency squeak production.

BACKGROUND: Anatomical and cognitive adaptations to overcome morpho-mechanical limitations of laryngeal sound production, where body size and the related vocal apparatus dimensions determine the fundamental frequency, increase vocal diversity across taxa. Elephants flexibly use laryngeal and trunk-based vocalizations to form a repertoire ranging from infrasonic rumbles to higher-pitched trumpets. Moreover, they are among the few evolutionarily distantly related animals (humans, pinnipeds, cetaceans, birds) capable of imitating species-atypical sounds. Yet, their vocal plasticity has so far not been related to functions within their natural communicative system, in part because not all call types have been systematically studied. Here, we reveal how Asian elephants (Elephas maximus) produce species-specific squeaks (F0 300-2300 Hz) by using acoustic camera recordings to visualize sound emission and examining this alongside acoustic, behavioral, and morphological data across seven captive groups. RESULTS: We found that squeaks were emitted through the closed mouth in synchrony with cheek depression and retraction of the labial angles. The simultaneous emission of squeaks with nasal snorts (biphonation) in one individual confirmed that squeak production was independent of nasal passage involvement and this implicated oral sound production. The squeaks' spectral structure is incongruent with laryngeal sound production and aerodynamic whistles, pointing to tissue vibration as the sound source. Anatomical considerations suggest that the longitudinal closed lips function as the vibrators. Acoustic and temporal parameters exhibit high intra- and inter-individual variability that enables individual but no call-subtype classification. Only 19 of 56 study subjects were recorded to squeak, mostly during alarming contexts and social arousal but some also on command. CONCLUSION: Our results strongly suggest that Asian elephants force air from the small oral cavity through the tensed lips, inducing self-sustained lip vibration. Besides human brass players, lip buzzing is not described elsewhere in the animal kingdom. Given the complexity of the proposed mechanism, the surprising absence of squeaking in most of the unrelated subjects and the indication for volitional control, we hypothesize that squeak production involves social learning. Our study offers new insights into how vocal and cognitive flexibility enables mammals to overcome size-related limitations of laryngeal sound production. This flexibility enables Asian elephants to exploit a frequency range spanning seven octaves within their communicative system.