This study used the source and filter theory approach to analyse sex differences in the acoustic features of African elephant (Loxodonta africana) 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 is partly, but not exclusively, responsible for 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.
Recent research reveals that giraffes (Giraffa camelopardalis sp.) exhibit a socially structured, fission–fusion system. In other species possessing this kind of society, information exchange is important and vocal communication is usually well developed. But is this true for giraffes? Giraffes are known to produce sounds, but there is no evidence that they use vocalizations for communication. Reports on giraffe vocalizations are mainly anecdotal and the missing acoustic descriptions make it difficult to establish a call nomenclature. Despite inconclusive evidence to date, it is widely assumed that giraffes produce infrasonic vocalizations similar to elephants. In order to initiate a more detailed investigation of the vocal communication in giraffes, we collected data of captive individuals during day and night. We particularly focussed on detecting tonal, infrasonic or sustained vocalizations. We collected over 947 h of audio material in three European zoos and quantified the spectral and temporal components of acoustic signals to obtain an accurate set of acoustic parameters. Besides the known burst, snorts and grunts, we detected harmonic, sustained and frequency-modulated “humming” vocalizations during night recordings. None of the recorded vocalizations were within the infrasonic range. These results show that giraffes do produce vocalizations, which, based on their acoustic structure, might have the potential to function as communicative signals to convey information about the physical and motivational attributes of the caller. The data further reveal that the assumption of infrasonic communication in giraffes needs to be considered with caution and requires further investigations in future studies.
6 September 2021 in Philosophical Transactions of the Royal Society B: Biological Sciences
Elephants exhibit remarkable vocal plasticity, and case studies reveal that individuals of African savannah (Loxodonta africana) and Asian (Elephas maximus) elephants are capable of vocal production learning. Surprisingly, however, little is known about contextual learning (usage and comprehension learning) in elephant communication. Usage learning can be demonstrated by training animals to vocalize in an arbitrary (cue-triggered) context. Here we show that adult African savannah elephants (n = 13) can vocalize in response to verbal cues, reliably producing social call types such as the low-frequency rumble, trumpets and snorts as well as atypical sounds using various mechanisms, thus displaying compound vocal control. We further show that rumbles emitted upon trainer cues differ significantly in structure from rumbles triggered by social contexts of the same individuals (n = 6). Every form of social learning increases the complexity of a communication system. In elephants, we only poorly understand their vocal learning abilities and the underlying cognitive mechanisms. Among other research, this calls for controlled learning experiments in which the prerequisite is operant/volitional control of vocalizations. This article is part of the theme issue ‘Vocal learning in animals and humans’.
How do elephants achieve their enormous vocal flexibility when communicating, imitating or creating idiosyncratic sounds? The mechanisms that underpin this trait combine motoric abilities with vocal learning processes. We demonstrate the unusual production techniques used by five African savanna elephants to create idiosyncratic sounds, which they learn to produce on cue by positive reinforcement training. The elephants generate these sounds by applying nasal tissue vibration via an ingressive airflow at the trunk tip, or by contracting defined superficial muscles at the trunk base. While the production mechanisms of the individuals performing the same sound categories are similar, they do vary in fine-tuning, revealing that each individual has its own specific sound-producing strategy. This plasticity reflects the creative and cognitive abilities associated with ‘vocal’ learning processes. The fact that these sounds were reinforced and cue-stimulated suggests that social feedback and positive reinforcement can facilitate vocal creativity and vocal learning behavior in elephants. Revealing the mechanism and the capacity for vocal learning and sound creativity is fundamental to understanding the eloquence within the elephants’ communication system. This also helps to understand the evolution of human language and of open-ended vocal systems, which build upon similar cognitive processes.
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.
Anthropogenic noise is an increasingly pervasive environmental stressor for wildlife, yet its behavioral impacts on large mammals remain poorly understood. Across the genus Giraffa, multiple giraffe species face varying degrees of conservation condern under IUCN criteria due to rapid habitat modification and increasing human disturbance, yet the behavioral consequences of anthropogenic noise exposure have not been experimentally tested. Because vigilance is a key anti‐predator response in large herbivores and is highly sensitive to disturbance, we used it as a focal metric to assess giraffe reactions to noise. We conducted multisite playback experiments across three reserves in the Free State Province, South Africa, that differed in human‐exposure levels. Free‐roaming giraffes were presented with three anthropogenic sound stimuli (drone, vehicle, people talking) and a natural control stimulus (ring‐necked dove). Vigilance responses were video‐recorded, quantified, and analyzed using mixed‐effects models. Giraffes at the low‐exposure site showed markedly stronger vigilance responses to anthropogenic than to the natural control, whereas responses at high‐exposure sites were weaker and less differentiated across stimuli. These site‐specific patterns indicate that ecological exposure history modulates giraffe responsiveness to noise. Across the full dataset, anthropogenic sounds consistently elicited stronger vigilance responses than the control stimulus, demonstrating that noise alone, independent of visual cues, can alter natural behavior in free‐roaming giraffes. Collectively, these findings highlight the importance of incorporating acoustic disturbance into environmental impact assessments and protected‐area management. More broadly, the results contribute to understanding how wildlife perceive and respond to human‐generated noise and emphasize the need to integrate soundscape considerations into conservation planning as anthropogenic noise continues to expand across savanna ecosystems.