Elephant Communication

A special issue of Animals (ISSN 2076-2615). This special issue belongs to the section "Wildlife".

Deadline for manuscript submissions: closed (31 January 2022) | Viewed by 57997

Special Issue Editors


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Guest Editor
Mammal Communication Lab, Department of Behavioural and Cognitive Biology, University of Vienna, Vienna, Austria
Interests: vocal communication; animal behavior; social learning; cognition

E-Mail Website
Guest Editor
Mammal Communication Lab, Department of Behavioural and Cognitive Biology, University of Vienna, Vienna, Austria
Interests: vocal communication; animal behavior; sexual communication

Special Issue Information

Dear Colleagues,

The living elephants are divided into three species within two genera, the African savannah (Loxodonta africana) and forest elephant (L. cyclotis) and the Asian elephant (Elephas maximus). Although living in substantially different habitats, they are all extremely social and cooperative, and intra-specific communication is highly developed. All social contacts are influenced by gestural, olfactory, tactile, vocal, and, potentially, seismic signals.

Elephants are distinct in various aspects. The proboscis, a muscular hydrostat and multifunctional organ, is multimodally used in social and reproductive events. Elephants have the most scent-capturing genes of any animal, and research has so far only scratched the surface of their world of olfaction. In the vocal domain, elephants are special due to their capacity for vocal production learning, a rare trait in non-human mammals. Elephants are long-lived, large brained, and cognitive animals, but we still know very little about how learning (i.e., social learning) affects elephant behavior in general.

Most research papers have focused on a single modality (e.g., sound or olfaction), but a more holistic approach is needed. This Special Issue aims to gather, in one publication, the most recent advances on elephant communication and social behavior. We welcome original papers, reviews, or case studies on Loxodonta spp. and Elephas maximus that reveal key characteristics of the species and genera. This Special Issue also hopes to stimulate discussion and collaboration among specialists and to establish ideas for further experimental research in order to assess the cognitive mechanisms underlying elephant communication and social behavior.

text

Dr. Angela S. Stoeger
Dr. Anton Baotic
Guest Editors

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Keywords

  • Elephants
  • communication
  • social behavior

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Published Papers (9 papers)

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Research

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22 pages, 3364 KiB  
Article
Sound Visualization Demonstrates Velopharyngeal Coupling and Complex Spectral Variability in Asian Elephants
by Veronika C. Beeck, Gunnar Heilmann, Michael Kerscher and Angela S. Stoeger
Animals 2022, 12(16), 2119; https://doi.org/10.3390/ani12162119 - 18 Aug 2022
Cited by 6 | Viewed by 4562
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 [...] Read more.
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. Full article
(This article belongs to the Special Issue Elephant Communication)
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23 pages, 18059 KiB  
Article
Male African Elephant (Loxodonta africana) Behavioral Responses to Estrous Call Playbacks May Inform Conservation Management Tools
by Caitlin E. O’Connell-Rodwell, Monica N. Sandri, Jodie L. Berezin, Jaquelyn M. Munevar, Colleen Kinzley, Jason D. Wood, Maggie Wiśniewska and J. Werner Kilian
Animals 2022, 12(9), 1162; https://doi.org/10.3390/ani12091162 - 1 May 2022
Cited by 2 | Viewed by 3611
Abstract
Driven by reproductive motives, male African elephants (Loxodonta africana) in musth often expand their home ranges to locate estrous females. This extended range, coupled with heightened aggression often observed in musth males, can be particularly problematic in regions where human-modified landscapes and [...] Read more.
Driven by reproductive motives, male African elephants (Loxodonta africana) in musth often expand their home ranges to locate estrous females. This extended range, coupled with heightened aggression often observed in musth males, can be particularly problematic in regions where human-modified landscapes and elephant territories increasingly overlap. Several mitigation tools have been tested to resolve a wide range of human–elephant conflicts with varying degrees of success due to geographical disparities and habituation. We present findings on the potential application of estrous call playbacks in manipulating the behavior and movement of male elephants non-invasively, particularly mature musth adults and younger post-dispersal males, in Etosha National Park. Estrous vocalizations were presented across 26 experimental trials to mature musth adults (n = 5), mature non-musth adults (n = 6), and non-musth males belonging to younger, post-dispersal age classes (n = 8), with behavioral responses scored on a gradient scale from 0–1. Both mature musth adults and younger non-musth elephants were significantly more likely to respond with the highest intensity by approaching the acoustic source compared to mature non-musth adults that avoided the call. However, younger males tested in the presence of an older, higher-ranking male tended to react with a lower intensity than those tested alone. This result likely demonstrates the influence of social hierarchy and associations on male elephant behavior. We also observed a significant increase in physiological response, measured by defecation rate, across all male groups in response to the estrous call playbacks. Our findings suggest that using estrous calls as acoustic deterrents may effectively and non-invasively aid in reducing tension at the human–elephant interface, depending on the age, social context, and reproductive status of the male elephant. Full article
(This article belongs to the Special Issue Elephant Communication)
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13 pages, 1232 KiB  
Article
Social Disruption Impairs Predatory Threat Assessment in African Elephants
by Graeme Shannon, Line S. Cordes, Rob Slotow, Cynthia Moss and Karen McComb
Animals 2022, 12(4), 495; https://doi.org/10.3390/ani12040495 - 17 Feb 2022
Cited by 7 | Viewed by 7006
Abstract
The transmission of reliable information between individuals is crucial for group-living animals. This is particularly the case for cognitively advanced mammals with overlapping generations that acquire detailed social and ecological knowledge over long lifetimes. Here, we directly compare the ecological knowledge of elephants [...] Read more.
The transmission of reliable information between individuals is crucial for group-living animals. This is particularly the case for cognitively advanced mammals with overlapping generations that acquire detailed social and ecological knowledge over long lifetimes. Here, we directly compare the ecological knowledge of elephants from two populations, with radically different developmental histories, to test whether profound social disruption affects their ability to assess predatory threat. Matriarchs (≤50 years of age) and their family groups received playbacks of three lions versus a single lion roaring. The family groups in the natural Amboseli population (Kenya) reliably assessed the greater predatory threat presented by three lions roaring versus one. However, in the socially disrupted Pilanesberg population (South Africa), no fine-scale distinctions were made between the numbers of roaring lions. Our results suggest that the removal of older and more experienced individuals in highly social species, such as elephants, is likely to impact the acquisition of ecological knowledge by younger group members, particularly through the lack of opportunity for social learning and cultural transmission of knowledge. This is likely to be exacerbated by the trauma experienced by juvenile elephants that witnessed the culling of family members and were translocated to new reserves. With increasing levels of anthropogenic disturbance, it is important that conservation practitioners consider the crucial role that population structure and knowledge transfer plays in the functioning and resilience of highly social and long-lived species. Full article
(This article belongs to the Special Issue Elephant Communication)
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13 pages, 3458 KiB  
Article
Social and Seasonal Factors Contribute to Shifts in Male African Elephant (Loxodonta africana) Foraging and Activity Patterns in Kruger National Park, South Africa
by Kara du Plessis, Stefanie Birgit Ganswindt, Henk Bertschinger, Bruce Crossey, Michelle Deborah Henley, Mmatsawela Ramahlo and André Ganswindt
Animals 2021, 11(11), 3070; https://doi.org/10.3390/ani11113070 - 27 Oct 2021
Cited by 3 | Viewed by 3627
Abstract
African savannah elephants (Loxodonta africana) are well-known as ecosystem engineers with the ability to modify vegetation structure. The present study aimed to examine how male elephant foraging behaviour is affected across (a) season (wet versus dry); (b) time of day (before [...] Read more.
African savannah elephants (Loxodonta africana) are well-known as ecosystem engineers with the ability to modify vegetation structure. The present study aimed to examine how male elephant foraging behaviour is affected across (a) season (wet versus dry); (b) time of day (before or after noon); (c) presence or absence of other elephants; and (d) reproductive state (musth versus no musth). Six radio-collared adult elephant bulls were observed twice per week from June 2007–June 2008 in Kruger National Park (KNP), South Africa. Using generalized linear mixed effect modeling, results indicate that elephant bulls graze more during the wet season and browse more during the dry season. To potentially offset the costs associated with thermoregulation during the heat of the day, KNP elephants spent more time foraging during the morning, and more time resting during the afternoon. Male elephants also foraged significantly less when they were associated with females compared to when they were alone or with other males. This is likely due to male–female associations formed mainly for reproductive purposes, thus impeding on male foraging behaviours. In contrast, the condition of musth, defined by the presence of related physical signs, had no significant effect on foraging behaviour. Full article
(This article belongs to the Special Issue Elephant Communication)
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15 pages, 599 KiB  
Article
Monitoring Behaviour in African Elephants during Introduction into a New Group: Differences between Related and Unrelated Animals
by Franziska Hörner, Ann-Kathrin Oerke, Dennis W. H. Müller, Uta Westerhüs, Idu Azogu-Sepe, Jiri Hruby and Gela Preisfeld
Animals 2021, 11(10), 2990; https://doi.org/10.3390/ani11102990 - 18 Oct 2021
Cited by 7 | Viewed by 4749
Abstract
The introduction of elephants into new groups is necessary for breeding programmes. However, behavioural studies on the reactions of these animals at first encounters are missing. In the present study, female African elephants (Loxodonta africana) living in zoos were observed during unifications with [...] Read more.
The introduction of elephants into new groups is necessary for breeding programmes. However, behavioural studies on the reactions of these animals at first encounters are missing. In the present study, female African elephants (Loxodonta africana) living in zoos were observed during unifications with unfamiliar elephants (introduction of two to one females and one to two females; n = 6) and reunifications with related elephants (two mother–daughter-pairs; n = 4) that were separated for 2 and 12 years, respectively. First encounters of the elephants were observed and recorded by scan sampling. The parameters measured were (a) signs of the characteristic Greeting Ceremony, (b) distance to the fence separating the elephants during first contact, and (c) time until trunks touched for the first time. The data were statistically analysed with SPSS. The results showed that related elephants performed a full Greeting Ceremony on reunifications. Unrelated elephants only expressed a minor greeting. During first encounters, related elephants predominantly showed affiliative behaviour (p = 0.001), whilst unrelated elephants expressed more agonistic behaviour (p = 0.001). The distance to the fence was significantly smaller for related elephants than for unrelated elephants (p = 0.038). first contact of trunks occurred on average after 3.00 s. in related elephants and 1026.25 s. in unrelated elephants. These findings indicate that related elephants recognise their kin after up to 12 years of separation, meet them with a full Greeting Ceremony during reunification, and seek contact to the related elephant, while unrelated elephants are hesitant during unifications with unfamiliar elephants and express more agonistic behaviour. The results testify that zoo elephants show the same species-specific social behaviour as their conspecifics in the wild. It also confirms the cognitive abilities of elephants and the significance of matrilines for breeding programmes. Full article
(This article belongs to the Special Issue Elephant Communication)
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18 pages, 1969 KiB  
Article
Function of Trunk-Mediated “Greeting” Behaviours between Male African Elephants: Insights from Choice of Partners
by Connie R. B. Allen, Darren P. Croft, Camille Testard and Lauren J. N. Brent
Animals 2021, 11(9), 2718; https://doi.org/10.3390/ani11092718 - 17 Sep 2021
Cited by 4 | Viewed by 4156
Abstract
A common behavioural interaction between male African elephants is for an actor to direct his trunk to contact a same sex conspecific’s mouth, temporal gland, or genital region. Such behaviours are often referred to as “greetings”. Along with its inherent tactile element, these [...] Read more.
A common behavioural interaction between male African elephants is for an actor to direct his trunk to contact a same sex conspecific’s mouth, temporal gland, or genital region. Such behaviours are often referred to as “greetings”. Along with its inherent tactile element, these behaviours also likely provide olfactory information to actors concerning aspects of the target’s phenotype, including sexual status, feeding history, individual identity, and emotional state. Here, we explore whether the age and novelty of potential interactors affect the choice of individuals targeted by male African elephants for these trunks to scent emitting organ (SEO) behaviours at social hotspots in a male-dominated area. Male elephants of all ages, except older adolescents aged 16–20 years, preferentially targeted elephants of the same age class for trunk-to-SEO behaviours. Elephants younger than 26 years did not direct trunk-to-SEO behaviours to mature bulls (26+ years) more than expected by chance, suggesting these behaviours are not primarily used for younger males to establish contact with, or obtain information from or about older, more experienced individuals. We also found no evidence that males directed these behaviours preferentially to new individuals they encountered at male aggregations (compared to those they arrived in groups with), suggesting these behaviours are not primarily employed by males as a reunion display to establish relationships between new individuals or update relationships between familiar individuals separated over time. Age-mates may be preferentially targeted with these behaviours as a means to facilitate further interaction with partners (e.g., for sparring activity), or as a safe way to assess relative dominance rank in similarly aged and hence, size and strength, matched dyads. Our results suggest male African elephants use close contact trunk-to-SEO behaviours continuously over time, to facilitate positive relationships, test willingness to interact, and assess aspects of phenotype, between males occupying the same ecological space. Full article
(This article belongs to the Special Issue Elephant Communication)
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Review

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17 pages, 1168 KiB  
Review
Acknowledging the Relevance of Elephant Sensory Perception to Human–Elephant Conflict Mitigation
by Robbie Ball, Sarah L. Jacobson, Matthew S. Rudolph, Miranda Trapani and Joshua M. Plotnik
Animals 2022, 12(8), 1018; https://doi.org/10.3390/ani12081018 - 14 Apr 2022
Cited by 8 | Viewed by 5972
Abstract
Elephants are well known for their socio-cognitive abilities and capacity for multi-modal sensory perception and communication. Their highly developed olfactory and acoustic senses provide them with a unique non-visual perspective of their physical and social worlds. The use of these complex sensory signals [...] Read more.
Elephants are well known for their socio-cognitive abilities and capacity for multi-modal sensory perception and communication. Their highly developed olfactory and acoustic senses provide them with a unique non-visual perspective of their physical and social worlds. The use of these complex sensory signals is important not only for communication between conspecifics, but also for decisions about foraging and navigation. These decisions have grown increasingly risky given the exponential increase in unpredictable anthropogenic change in elephants’ natural habitats. Risk taking often develops from the overlap of human and elephant habitat in Asian and African range countries, where elephants forage for food in human habitat and crop fields, leading to conflict over high-quality resources. To mitigate this conflict, a better understanding of the elephants’ sensory world and its impact on their decision-making process should be considered seriously in the development of long-term strategies for promoting coexistence between humans and elephants. In this review, we explore the elephants’ sensory systems for audition and olfaction, their multi-modal capacities for communication, and the anthropogenic changes that are affecting their behavior, as well as the need for greater consideration of elephant behavior in elephant conservation efforts. Full article
(This article belongs to the Special Issue Elephant Communication)
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21 pages, 2771 KiB  
Review
Does Social Complexity Drive Vocal Complexity? Insights from the Two African Elephant Species
by Daniela Hedwig, Joyce Poole and Petter Granli
Animals 2021, 11(11), 3071; https://doi.org/10.3390/ani11113071 - 27 Oct 2021
Cited by 11 | Viewed by 6144
Abstract
The social complexity hypothesis (SCH) for communication states that the range and frequency of social interactions drive the evolution of complex communication systems. Surprisingly, few studies have empirically tested the SHC for vocal communication systems. Filling this gap is important because a co-evolutionary [...] Read more.
The social complexity hypothesis (SCH) for communication states that the range and frequency of social interactions drive the evolution of complex communication systems. Surprisingly, few studies have empirically tested the SHC for vocal communication systems. Filling this gap is important because a co-evolutionary runaway process between social and vocal complexity may have shaped the most intricate communication system, human language. We here propose the African elephant Loxodonta spec. as an excellent study system to investigate the relationships between social and vocal complexity. We review how the distinct differences in social complexity between the two species of African elephants, the forest elephant L. cyclotis and the savanna elephant L. africana, relate to repertoire size and structure, as well as complex communication skills in the two species, such as call combination or intentional formant modulation including the trunk. Our findings suggest that Loxodonta may contradict the SCH, as well as other factors put forth to explain patterns of vocal complexity across species. We propose that life history traits, a factor that has gained little attention as a driver of vocal complexity, and the extensive parental care associated with a uniquely low and slow reproductive rate, may have led to the emergence of pronounced vocal complexity in the forest elephant despite their less complex social system compared to the savanna elephant. Conclusions must be drawn cautiously, however. A better understanding of vocal complexity in the genus Loxodonta will depend on continuing advancements in remote data collection technologies to overcome the challenges of observing forest elephants in their dense rainforest habitat, as well as the availability of directly comparable data and methods, quantifying both structural and contextual variability in the production of rumbles and other vocalizations in both species of African elephants. Full article
(This article belongs to the Special Issue Elephant Communication)
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18 pages, 3737 KiB  
Review
The Chemical Ecology of Elephants: 21st Century Additions to Our Understanding and Future Outlooks
by Bruce A. Schulte and Chase A. LaDue
Animals 2021, 11(10), 2860; https://doi.org/10.3390/ani11102860 - 30 Sep 2021
Cited by 13 | Viewed by 12874
Abstract
Chemical signals are the oldest and most ubiquitous means of mediating intra- and interspecific interactions. The three extant species of elephants, the Asian elephant and the two African species, savanna and forest share sociobiological patterns in which chemical signals play a vital role. [...] Read more.
Chemical signals are the oldest and most ubiquitous means of mediating intra- and interspecific interactions. The three extant species of elephants, the Asian elephant and the two African species, savanna and forest share sociobiological patterns in which chemical signals play a vital role. Elephants emit secretions and excretions and display behaviors that reveal the importance of odors in their interactions. In this review, we begin with a brief introduction of research in elephant chemical ecology leading up to the 21st century, and then we summarize the body of work that has built upon it and occurred in the last c. 20 years. The 21st century has expanded our understanding on elephant chemical ecology, revealing their use of odors to detect potential threats and make dietary choices. Furthermore, complementary in situ and ex situ studies have allowed the careful observations of captive elephants to be extended to fieldwork involving their wild counterparts. While important advances have been made in the 21st century, further work should investigate the roles of chemical signaling in elephants and how these signals interact with other sensory modalities. All three elephant species are threatened with extinction, and we suggest that chemical ecology can be applied for targeted conservation efforts. Full article
(This article belongs to the Special Issue Elephant Communication)
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