Field Propagation Experiments of Male African Savanna Elephant Rumbles: A Focus on the Transmission of Formant Frequencies.

African savanna elephants live in dynamic fission⁻fusion societies and exhibit a sophisticated vocal communication system. Their most frequent call-type is the 'rumble', with a fundamental frequency (which refers to the lowest vocal fold vibration rate when producing a vocalization) near or in the infrasonic range. Rumbles are used in a wide variety of behavioral contexts, for short- and long-distance communication, and convey contextual and physical information. For example, maturity (age and size) is encoded in male rumbles by formant frequencies (the resonance frequencies of the vocal tract), having the most informative power. As sound propagates, however, its spectral and temporal structures degrade progressively. Our study used manipulated and resynthesized male social rumbles to simulate large and small individuals (based on different formant values) to quantify whether this phenotypic information efficiently transmits over long distances. To examine transmission efficiency and the potential influences of ecological factors, we broadcasted and re-recorded rumbles at distances of up to 1.5 km in two different habitats at the Addo Elephant National Park, South Africa. Our results show that rumbles were affected by spectral⁻temporal degradation over distance. Interestingly and unlike previous findings, the transmission of formants was better than that of the fundamental frequency. Our findings demonstrate the importance of formant frequencies for the efficiency of rumble propagation and the transmission of information content in a savanna elephant's natural habitat

MANOVA comparing source- and filter-related acoustic features between male and female African elephant social rumbles.

<p>MANOVA comparing source- and filter-related acoustic features between male and female African elephant social rumbles.</p

Sexual dimorphism in African elephant social rumbles

<div><p>This study used the source and filter theory approach to analyse sex differences in the acoustic features of African elephant (<i>Loxodonta africana</i>) low-frequency rumbles produced in social contexts (‘social rumbles’). Permuted discriminant function analysis revealed that rumbles contain sufficient acoustic information to predict the sex of a vocalizing individual. Features primarily related to the vocalizer’s size, i.e. fundamental frequency variables and vocal tract resonant frequencies, differed significantly between the sexes. Yet, controlling for age and size effects, our results indicate that the pronounced sexual size dimorphism in African elephants <i>is partly</i>, <i>but not exclusively</i>, <i>responsible for</i> sexual differences in social rumbles. This provides a scientific foundation for future work investigating the perceptual and functional relevance of specific acoustic characteristics in African elephant vocal sexual communication.</p></div

Field Propagation Experiments of Male African Savanna Elephant Rumbles: A Focus on the Transmission of Formant Frequencies

African savanna elephants live in dynamic fission&ndash;fusion societies and exhibit a sophisticated vocal communication system. Their most frequent call-type is the &lsquo;rumble&rsquo;, with a fundamental frequency (which refers to the lowest vocal fold vibration rate when producing a vocalization) near or in the infrasonic range. Rumbles are used in a wide variety of behavioral contexts, for short- and long-distance communication, and convey contextual and physical information. For example, maturity (age and size) is encoded in male rumbles by formant frequencies (the resonance frequencies of the vocal tract), having the most informative power. As sound propagates, however, its spectral and temporal structures degrade progressively. Our study used manipulated and resynthesized male social rumbles to simulate large and small individuals (based on different formant values) to quantify whether this phenotypic information efficiently transmits over long distances. To examine transmission efficiency and the potential influences of ecological factors, we broadcasted and re-recorded rumbles at distances of up to 1.5 km in two different habitats at the Addo Elephant National Park, South Africa. Our results show that rumbles were affected by spectral&ndash;temporal degradation over distance. Interestingly and unlike previous findings, the transmission of formants was better than that of the fundamental frequency. Our findings demonstrate the importance of formant frequencies for the efficiency of rumble propagation and the transmission of information content in a savanna elephant&rsquo;s natural habitat

Comparison of a female (left, Chikwenya) and a male (right, Mike) African elephant.

<p>The spectrogram and power spectra below the photograph provide an example of a social rumble of each sex, indicating formant positions (rumbles uttered first by Chikwenya, followed by Mike).</p

Study sites, sex, age and number of calls for each study subject.

<p>Study sites, sex, age and number of calls for each study subject.</p

The vocal repertoire of infant giant pandas (<i>Ailuropoda melanoleuca</i>)

<p>Infant giant pandas (<i>Ailuropoda melanoleuca</i>) are highly vocal during the first few weeks of their life. Despite this, no previous studies have attempted to systematically categorize infant giant panda vocalizations into different call types. In this study, we used acoustic and video analyses to split infant giant panda vocalizations into three distinct call types based on their acoustic structure as well as their use in different behavioural contexts. A discriminant functions analysis on the acoustic variables confirmed our initial subjective classification of 281 vocalizations into three call types: the harsh sounding “squawk”, the high-pitched “squall” and the pulsed “croak”. Based on the observed spectral acoustic characteristics, none of these three infant call types appears to be a precursor of an adult giant panda vocalization. In addition, individual call types could not be assigned to specific recording contexts. These findings suggest that infant giant panda vocalizations convey information about a cub's distress and need, rather than being tied to specific contexts of emission. Our objective demonstration that infant giant pandas have three basic call types provides a foundation for future studies of vocal ontogeny in this highly endangered species.</p

Field Propagation Experiments of Male African Savanna Elephant Rumbles: A Focus on the Transmission of Formant Frequencies

African savanna elephants live in dynamic fission–fusion societies and exhibit a sophisticated vocal communication system. Their most frequent call-type is the ‘rumble’, with a fundamental frequency (which refers to the lowest vocal fold vibration rate when producing a vocalization) near or in the infrasonic range. Rumbles are used in a wide variety of behavioral contexts, for short- and long-distance communication, and convey contextual and physical information. For example, maturity (age and size) is encoded in male rumbles by formant frequencies (the resonance frequencies of the vocal tract), having the most informative power. As sound propagates, however, its spectral and temporal structures degrade progressively. Our study used manipulated and resynthesized male social rumbles to simulate large and small individuals (based on different formant values) to quantify whether this phenotypic information efficiently transmits over long distances. To examine transmission efficiency and the potential influences of ecological factors, we broadcasted and re-recorded rumbles at distances of up to 1.5 km in two different habitats at the Addo Elephant National Park, South Africa. Our results show that rumbles were affected by spectral–temporal degradation over distance. Interestingly and unlike previous findings, the transmission of formants was better than that of the fundamental frequency. Our findings demonstrate the importance of formant frequencies for the efficiency of rumble propagation and the transmission of information content in a savanna elephant’s natural habitat.© 2018 by the author

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

Male African elephants discriminate and prefer vocalizations of unfamiliar females

Gaining information about conspecifics via long-distance vocalizations is crucial for social and spatially flexible species such as the African elephant (Loxodonta africana). Female elephants are known to discriminate individuals and kin based on acoustic cues. Specifically, females approached the loudspeaker exclusively with playbacks of familiar individuals with high association indexes, intentionally fusing with their affiliates. For males, which are less bonded, gathering social information via vocalizations could still have important implications, but little is known about their vocal discrimination skills. We experimentally tested the ability of male African elephants to discriminate the social rumbles of familiar (from the same population) versus unfamiliar females. Male elephants discriminated and preferentially moved towards the rumbles of unfamiliar females, showing longer attentive reactions and significantly more orientating (facing and approaching the speaker) behavior. The increased orientating response of males towards playbacks of unfamiliar females is converse to the reaction of female subjects. Our results provide evidence that male elephants extract social information from vocalizations, yet with a different intention than females. Accordingly, males might use social cues in vocalizations to assess mating opportunities, which may involve selection to identify individuals or kin in order to avoid inbreeding.© The Author(s) 201