Angela S. Stoeger

How Low Can You Go? Physical Production Mechanism of Elephant Infrasonic Vocalizations

The Song of the Elephant In mammals, vocal sound production generally occurs in one of two ways, either through muscular control—as when a cat purrs or, more commonly, by air passing through the vocal folds—which occurs in humans and facilitates production of extremely high frequency bat calls. Over the past 20 years, it has been recognized that elephants can communicate through extremely low frequency infrasonic sounds. Taking advantage of a natural death of an elephant in a zoo, Herbst et al. (p. 595) examined the biomechanics of elephant sound production in an excised elephant larynx. Self-sustained vocal-fold vibrations, without the presence of any neural control, were used to produce infrasonic elephant sounds, using the same mechanism as singing in humans and echolocation in bats. Elephants produce low-frequency sounds via intrinsic vocal-fold vibrations similar to those in humans. Elephants can communicate using sounds below the range of human hearing (“infrasounds” below 20 hertz). It is commonly speculated that these vocalizations are produced in the larynx, either by neurally controlled muscle twitching (as in cat purring) or by flow-induced self-sustained vibrations of the vocal folds (as in human speech and song). We used direct high-speed video observations of an excised elephant larynx to demonstrate flow-induced self-sustained vocal fold vibration in the absence of any neural signals, thus excluding the need for any “purring” mechanism. The observed physical principles of voice production apply to a wide variety of mammals, extending across a remarkably large range of fundamental frequencies and body sizes, spanning more than five orders of magnitude.

An Asian elephant imitates human speech.

Summary Vocal imitation has convergently evolved in many species, allowing learning and cultural transmission of complex, conspecific sounds, as in birdsong [1, 2]. Scattered instances also exist of vocal imitation across species, including mockingbirds imitating other species or parrots and mynahs producing human speech [3, 4]. Here, we document a male Asian elephant (Elephas maximus) that imitates human speech, matching Korean formants and fundamental frequency in such detail that Korean native speakers can readily understand and transcribe the imitations. To create these very accurate imitations of speech formant frequencies, this elephant (named Koshik) places his trunk inside his mouth, modulating the shape of the vocal tract during controlled phonation. This represents a wholly novel method of vocal production and formant control in this or any other species. One hypothesized role for vocal imitation is to facilitate vocal recognition by heightening the similarity between related or socially affiliated individuals [1, 2]. The social circumstances under which Koshik’s speech imitations developed suggest that one function of vocal learning might be to cement social bonds and, in unusual cases, social bonds across species.

Vocal cues indicate level of arousal in infant African elephant roars

Arousal-based physiological changes influence acoustic features of vocalizations in mammals. In particular, nonlinear phenomena are thought to convey information about the callers arousal state. This hypothesis was tested in the infant African elephant (Loxodonta africana) roar, a call type produced in situations of arousal and distress. Ninety-two percent of roars exhibited nonlinear phenomena, with chaos being the most common type. Acoustic irregularities were strongly associated with elevated fundamental frequency values. Roars produced in situations of highest urgency, based on the occurrence of behavioral indicators of arousal, were characterized by the lowest harmonics-to-noise ratio; this indicates low tonality. In addition, roars produced in these situations lasted longer than those produced in contexts of lower presumed urgency. Testing the infant roars for individual distinctiveness revealed only a moderate classification result. Combined, these findings indicate that infant African elephant roars primarily function to signal the callers arousal state. The effective communication of this type of information may allow mothers to respond differentially based on their infants degree of need and may be crucial for the survival of infant African elephants in their natural environment.

Vocal learning in elephants: neural bases and adaptive context.

In the last decade clear evidence has accumulated that elephants are capable of vocal production learning. Examples of vocal imitation are documented in African (Loxodonta africana) and Asian (Elephas maximus) elephants, but little is known about the function of vocal learning within the natural communication systems of either species. We are also just starting to identify the neural basis of elephant vocalizations. The African elephant diencephalon and brainstem possess specializations related to aspects of neural information processing in the motor system (affecting the timing and learning of trunk movements) and the auditory and vocalization system. Comparative interdisciplinary (from behavioral to neuroanatomical) studies are strongly warranted to increase our understanding of both vocal learning and vocal behavior in elephants.

Towards an automated acoustic detection system for free-ranging elephants

The human–elephant conflict is one of the most serious conservation problems in Asia and Africa today. The involuntary confrontation of humans and elephants claims the lives of many animals and humans every year. A promising approach to alleviate this conflict is the development of an acoustic early warning system. Such a system requires the robust automated detection of elephant vocalizations under unconstrained field conditions. Today, no system exists that fulfils these requirements. In this paper, we present a method for the automated detection of elephant vocalizations that is robust to the diverse noise sources present in the field. We evaluate the method on a data-set recorded under natural field conditions to simulate a real-world scenario. The proposed method outperformed existing approaches and robustly and accurately detected elephants. It thus can form the basis for a future automated early warning system for elephants. Furthermore, the method may be a useful tool for scientists in bioacoustics for the study of wildlife recordings.

Age-group estimation in free-ranging African elephants based on acoustic cues of low-frequency rumbles

Animal vocal signals are increasingly used to monitor wildlife populations and to obtain estimates of species occurrence and abundance. In future, acoustic monitoring should function not only to detect animals, but also to extract detailed information about populations by discriminating sexes, age groups, social or kin groups, and potentially individuals. Here we show that it is possible to estimate age groups of African elephants (Loxodonta africana) based on acoustic parameters extracted from rumbles recorded under field conditions in a National Park in South Africa. Statistical models reached up to 70% correct classification to four age groups (infants, calves, juveniles and adults) and 95% correct classification when categorizing into two groups (infants/calves lumped into one group vs. adults). The models revealed that parameters representing absolute frequency values have the most discriminative power. Comparable classification results were obtained by fully automated classification of rumbles by high-dimensional features that represent the entire spectral envelope, such as Mel-frequency cepstral coefficient (75% correct classification) and Greenwood function cepstral coefficient (74% correct classification). The reported results and methods provide the scientific foundation for a future system that could potentially automatically estimate the demography of an acoustically monitored elephant group or population.

Establishing the fundamentals for an elephant early warning and monitoring system

BackgroundThe decline of habitat for elephants due to expanding human activity is a serious conservation problem. This has continuously escalated the human–elephant conflict in Africa and Asia. Elephants make extensive use of powerful infrasonic calls (rumbles) that travel distances of up to several kilometers. This makes elephants well-suited for acoustic monitoring because it enables detecting elephants even if they are out of sight. In sight, their distinct visual appearance makes them a good candidate for visual monitoring. We provide an integrated overview of our interdisciplinary project that established the scientific fundamentals for a future early warning and monitoring system for humans who regularly experience serious conflict with elephants. We first draw the big picture of an early warning and monitoring system, then review the developed solutions for automatic acoustic and visual detection, discuss specific challenges and present open future work necessary to build a robust and reliable early warning and monitoring system that is able to operate in situ.FindingsWe present a method for the automated detection of elephant rumbles that is robust to the diverse noise sources present in situ. We evaluated the method on an extensive set of audio data recorded under natural field conditions. Results show that the proposed method outperforms existing approaches and accurately detects elephant rumbles. Our visual detection method shows that tracking elephants in wildlife videos (of different sizes and postures) is feasible and particularly robust at near distances.DiscussionFrom our project results we draw a number of conclusions that are discussed and summarized. We clearly identified the most critical challenges and necessary improvements of the proposed detection methods and conclude that our findings have the potential to form the basis for a future automated early warning system for elephants. We discuss challenges that need to be solved and summarize open topics in the context of a future early warning and monitoring system. We conclude that a long-term evaluation of the presented methods in situ using real-time prototypes is the most important next step to transfer the developed methods into practical implementation.

Information content and acoustic structure of male African elephant social rumbles.

Until recently, the prevailing theory about male African elephants (Loxodonta africana) was that, once adult and sexually mature, males are solitary and targeted only at finding estrous females. While this is true during the state of ‘musth’ (a condition characterized by aggressive behavior and elevated androgen levels), ‘non-musth’ males exhibit a social system seemingly based on companionship, dominance and established hierarchies. Research on elephant vocal communication has so far focused on females, and very little is known about the acoustic structure and the information content of male vocalizations. Using the source and filter theory approach, we analyzed social rumbles of 10 male African elephants. Our results reveal that male rumbles encode information about individuality and maturity (age and size), with formant frequencies and absolute fundamental frequency values having the most informative power. This first comprehensive study on male elephant vocalizations gives important indications on their potential functional relevance for male-male and male-female communication. Our results suggest that, similar to the highly social females, future research on male elephant vocal behavior will reveal a complex communication system in which social knowledge, companionship, hierarchy, reproductive competition and the need to communicate over long distances play key roles.

Vocal stress associated with a translocation of a family herd of African elephants (Loxodonta africana) in the Kruger National Park, South Africa

We used vocal indicators to examine the effect of a translocation of an African elephant family herd within the Kruger National Park. These animals were moved 300 km from their home range, but returned unaided to this range within 23 days. We found that translocation resulted in a change in the mean fundamental frequency of low-frequency elephant vocalizations, known as rumbles. The rumbles increased significantly in pitch compared with pre-translocation levels during the 23 days the animals spent outside their normal home range. Mean fundamental frequency returned close to pre-translocation level by the time the animals had navigated their way back to their previous home range. Raised pitch is known to be an indicator of stress in humans and other animals. The observed acoustic results are consistent with a physiological measure of stress, faecal glucocorticoid metabolite levels, which were monitored from the same animals during the study and have already been reported elsewhere. To our knowledge, this is the first report of prolonged monitoring of vocal stress response in free-ranging animals. Measuring behavioural responses, such as vocalizations, may provide an objective non-invasive method for assessing stress. This could help in determining the effects that particular management actions might have on elephants.

Vocalizations and associated behaviour of Asian elephant (Elephas maximus) calves

The present study describes the acoustic structure and associated behavioural contexts of vocalizations of Asian elephant calves. Three hundred twenty-seven vocalizations of six calves aged between 6 and 27 months were recorded at the Emmen and the Cologne zoo. Based on temporal and structural characteristics including nonlinear phenomena, we structurally distinguished four call types, the rumble, the roar, the chirp and the trumpet. Most vocalizations were uttered during all forms of play behaviour, with chirps and trumpets dominating here. Rumbles were mainly recorded in combination with roars; the latter were highly chaotic in structure and associated with higher arousal contexts (e.g., being separated from the mother). These first insights into the vocal behaviour of Asian elephant calves reveal that calf vocalizations clearly differ in acoustic structure and application from those of adults and older juveniles. This highlights the importance of investigating the vocal ontogeny of elephants as well as other species having a long adolescence and developmental period.