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

SIMPLE SUMMARY: Whether a vocalization is emitted though the mouth or nose impacts on its acoustic shape through specific vocal tract resonances. In human language, nasalized compared to oral vowels can change the meaning of words. African elephants mainly use low-frequency rumbles to communicate and can utter them either through the mouth or trunk. In this study, we used an acoustic camera to visualize the sound emission of rumbles in Asian elephants, which have been studied considerably less than African elephants. We recorded nine captive female Asian elephants and analyzed the acoustic structure of 203 calls. We found that most rumbles (64%) were emitted through the trunk, 21% through the mouth, and surprisingly, 13% simultaneously through the mouth and trunk. Some of the rumbles were combined with orally emitted roars. The nasal rumbles concentrated most spectral energy in lower frequencies, whereas the oral and mixed rumbles showed a broader spectral energy distribution and concentration in higher frequencies, and were louder. The roars were the loudest, broadest and highest in frequency. This study is the first to demonstrate coupled oral and nasal emission in a non–human animal, thereby setting an important framework to study the functions of this acoustic variability in elephant communication and the evolution of vocal flexibility across species. ABSTRACT: 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.