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Elephant behavior toward the dead: A review and insights from field observations
Abstract
Many nonhuman animals have been documented to take an interest in their dead. A few socially complex and cognitively advanced taxa—primates, cetaceans, and proboscideans—stand out for the range and duration of behaviors that they display at conspecific carcasses. Here, we review the literature on field observations of elephants at carcasses to identify patterns in behaviors exhibited. We add to this literature by describing elephant responses to dead elephants in the Samburu National Reserve, northern Kenya. The literature review indicated that behavior of elephants at carcasses most often included approaches, touching, and investigative responses, and these occurred at varying stages of decay, from fresh carcasses to scattered and sun-bleached bones. During our own observations, we also witnessed elephants visiting and revisiting carcasses during which they engaged in extensive investigative behavior, stationary behavior, self-directed behavior, temporal gland streaming, and heightened social interactions with other elephants in the vicinity of a carcass. Elephants show broad interest in their dead regardless of the strength of former relationships with the dead individual. Such behaviors may allow them to update information regarding their social context in this highly fluid fission–fusion society. The apparent emotionality and widely reported inter-individual differences involved in elephant responses to the dead deserve further study. Our research contributes to the growing discipline of comparative thanatology to illuminate the cognition and context of nonhuman animal response to death, particularly among socially complex species.
... Most animal species, unlike humans, pay less interest to their dead conspecifics. Different species' 'weak' cognitive abilities and disease avoidance theories support this 'leaving alone' behaviour (Goldenberg & Wittemyer, 2019). However, cetaceans, primates, and elephants show contrasting examples of strong behavioural reactions to their deceased young ones. ...
... For instance, a behaviour where the elephant carcass parts were carried across a distance, covering the carcass with vegetation ("weak" burial), or observing the carcass over several hours has been covered in various scholarships concerning African elephants (Douglas-Hamilton and Douglas-Hamilton 1975;Goldenberg & Wittemyer, 2019). Apart from elephants, burial behaviour has been observed in termites (Reticulitermes fukienensis ) (Fernández-fueyo et al. 2021) but in no other large mammals. ...
... Calf burials by elephants are among the less-studied topics of thanatology, especially in Asia (Sharma et al. 2019), even though this practice is known to the conservation gatekeepers. Elephants show a variety of behaviour ranging from investigative, stationary, self-directed, social, and mourning behaviour around their dead conspecific (Hawley et al. 2016;Goldenberg and Wittemyer 2019;Watts 2019;Stephan et al. 2020). ...
Rampant environmental change and forest destruction push elephants, both Asian and African, to explore human spaces to fulfil their dietary and ecological requirements. Many ‘novel’ elephant behaviour in sharing spaces come to the limelight. Elephant calf burial is reported in African literature but remains absent from the Asian context. We concretely report calf burials by Asian elephants in the eastern Himalayan floodplains of the north Bengal landscape. The study area consists of fragmented forests, tea gardens, agricultural lands, and defence establishments, among others. Tea gardens form the majority of elephant corridors, and we explain the burial strategy of elephants in the trenches of tea gardens. We present four case reports of calf burials by elephants. We aimed to understand the perimortem strategy and postmortem behaviour of Asian elephants. The major findings reflect that the carcasses were carried through trunks and legs for a distance before burying in a ‘legs-upright-position’. We further investigated the underlying reason for calf deaths through postmortem examinations. Direct human intervention was not recorded in any of the four deaths. Through opportunistic observation, digital photography and fieldnotes, and postmortem examination report, we suggest that the carcasses were buried in an abnormal recumbent style irrespective of the reason for calf deaths. Through long-term observation, we further report that the elephants in this region clearly avoid the paths where carcasses were buried, attributing to “bad milestones” and “bad omens”. We discuss and connect the literature of two distinct elephant species and also compared thanatological studies of other sentient nonhuman species. Keywords: Thanatology, animal behaviour, Asian elephant, calf burial, eastern Himalayas, tea gardens
... Some mammalian species, such as proboscideans, that live in complex societies with strong social bonds among kin and possessing higher cognitive abilities perhaps display greater interest towards their dead conspecifics. In the African savanna elephant (Loxodonta africana), several studies have documented strong interest in dead conspecifics (expressed by investigating, repeatedly visiting or touching the deceased, as well as self-directed reactions), irrespective of their genetic relationship with the dead individual [22][23][24][25]. They show greater inquisitive behaviour towards deceased conspecifics than heterospecifics [26]. ...
... In the absence of confirmed evidence on relatedness, we may only speculate that the carcasses were carried by their mothers. Similar behaviour has also been observed in African savanna elephants, who were known to carry pieces of jaws, rib bones and even full-grown tusks (of dead adults), from short to long distances [16,22,25,48,49]. Carrying and interacting with fragments of carcasses are yet to be observed in Asian elephants. ...
... Other touch responses included trunk rest, tapping and shaking the trunk or leg of the deceased, and climbing on and mounting-like postures on the dead body (by young individuals). Touching the corpse was also the most frequently observed behaviour in African elephants [25]. These observations highlight the importance of tactile communication in elephant societies. ...
Documenting the behavioural repertoire of an animal species is important for understanding that species' natural history. Many behaviours such as mating, parturition and death may be observed only rarely in the wild due to the low frequency of occurrence, short duration and the species' elusiveness. Opportunistic documentation of rare behaviours is therefore valuable for deciphering the behavioural complexity in a species. In this context, digital platforms may serve as useful data sources for studying rare behaviours in animals. Using videos uploaded on YouTube, we document and construct a tentative repertoire of thanatological responses (death-related behaviours) in Asian elephants ( Elephas maximus ). The most frequently observed thanatological responses included postural changes, guarding/keeping vigil, touching, investigating the carcass, epimeletic behaviours and vocalizations. We also describe some infrequently observed behaviours, including carrying dead calves by adult females, re-assurance-like behaviours and attempts to support dying or dead conspecifics, some of which were only known anecdotally in Asian elephants. Our observations indicate the significance of open-source video data on digital platforms for gaining insights into rarely observed behaviours and support the accumulating evidence for higher cognitive abilities of Asian elephants in the context of comparative thanatology.
... Elephant post-mortem attentive behaviour has been documented in all three extant species of elephants [6,-8]. Long known to elephant researchers, interest in dead conspecifics is not only limited to carcases but also extends towards conspecific bones and tusks [9]. During a study of elephant carcase decomposition, the skull was transported as far as 100 m from the original site by other elephants [10]. ...
... The results showed that elephants (i) approached and manipulated elephant tusks significantly more than other objects, (ii) similarly showed more interest in conspecific skulls than non-conspecific skulls, and (iii) appeared not to differentiate skulls of previously known individuals from the skulls of strangers. Their main findings are further supported by an informal experiment conducted by Goldenberg and Wittemyer [9] in which they presented elephant, giraffe and Cape buffalo bones to wild elephants, with the most interest shown to elephant bones. Similar results were obtained in captive elephants (Rassmussen in [9]). ...
... Their main findings are further supported by an informal experiment conducted by Goldenberg and Wittemyer [9] in which they presented elephant, giraffe and Cape buffalo bones to wild elephants, with the most interest shown to elephant bones. Similar results were obtained in captive elephants (Rassmussen in [9]). ...
Chimpanzees exhibit a variety of behaviours surrounding their dead, although much less is known about how they respond towards conspecific skeletons. We tested chimpanzees' visual attention to images of conspecific and non-conspecific stimuli (cat/chimp/dog/rat), shown simultaneously in four corners of a screen in distinct orientations (frontal/diagonal/lateral) of either one of three types (faces/skulls/skull-shaped stones). Additionally, we compared their visual attention towards chimpanzee-only stimuli (faces/skulls/skull-shaped stones). Lastly, we tested their attention towards specific regions of chimpanzee skulls. We theorized that chimpanzee skulls retaining face-like features would be perceived similarly to chimpanzee faces and thus be subjected to similar biases. Overall, supporting our hypotheses, the chimpanzees preferred conspecific-related stimuli. The results showed that chimpanzees attended: (i) significantly longer towards conspecific skulls than other species skulls (particularly in forward-facing and to a lesser extent diagonal orientations); (ii) significantly longer towards conspecific faces than other species faces at forward-facing and diagonal orientations; (iii) longer towards chimpanzee faces compared with chimpanzee skulls and skull-shaped stones, and (iv) attended significantly longer to the teeth, similar to findings for elephants. We suggest that chimpanzee skulls retain relevant, face-like features that arguably activate a domain-specific face module in chimpanzees' brains, guiding their attention.
... It is widely accepted that elephants have an awareness of and a general interest in death (Sharma et al. 2019). Male elephants approach the carcasses of other males as a form of dominance, obtaining necessary information about the deceased and engaging in displacement behaviors; female herds have been observed to visit the carcasses of unrelated matriarchs or multiparous females, directly investigating the carcass (Douglas-Hamilton et al. 2006;Merte et al. 2009;Hawley et al. 2018;Goldenberg & Wittemyer 2019). Such interest is not merely limited to kin; African elephants have been observed to direct attention toward dead conspecifics regardless of relationship strength with the deceased (Goldenberg & Wittemyer 2019). ...
... Male elephants approach the carcasses of other males as a form of dominance, obtaining necessary information about the deceased and engaging in displacement behaviors; female herds have been observed to visit the carcasses of unrelated matriarchs or multiparous females, directly investigating the carcass (Douglas-Hamilton et al. 2006;Merte et al. 2009;Hawley et al. 2018;Goldenberg & Wittemyer 2019). Such interest is not merely limited to kin; African elephants have been observed to direct attention toward dead conspecifics regardless of relationship strength with the deceased (Goldenberg & Wittemyer 2019). However, it is unknown as to whether there is a difference between a carcass and bones; their extensive investigation of a carcass, occurring at various stages of carcass decay, including sniffing the deceased is seen to represent processing of death (Hawley et al. 2018;Goldenberg & Wittemyer 2019). ...
... Such interest is not merely limited to kin; African elephants have been observed to direct attention toward dead conspecifics regardless of relationship strength with the deceased (Goldenberg & Wittemyer 2019). However, it is unknown as to whether there is a difference between a carcass and bones; their extensive investigation of a carcass, occurring at various stages of carcass decay, including sniffing the deceased is seen to represent processing of death (Hawley et al. 2018;Goldenberg & Wittemyer 2019). This then leads one to question whether without the visual and/or olfactory cues that surround a carcass, an individual would recognize death? ...
Elephants are highly social beings with complex individual personalities. We know that elephants have a general interest in death, investigating carcasses, not just limited to kin; however, research does not explore in depth whether individuals change their behavior or personality following traumatic events, such as the death of a conspecific. Within a captive herd of Asian elephants (Elephas maximus ) housed at Chester Zoo, UK, we measured social behaviour and proximity and personality using the TIPI, and found age‐related and relationship‐related changes in both behavior and personality following the deaths of herd members. Overall, the herd spent less time socialising and engaging in affiliative behaviors following the death of the adult female when compared to baseline data, yet spent more time engaging in these behaviors after the death of two calves. The death of the central female had a dramatic impact on her infant calf, resulting in increasingly withdrawn behavior, yet had the opposite effect on her adult daughter, who subsequently established a more integrated role within the herd. Emotional Stability fell in the motherless calf but rose in an adult female, who had lost her adult daughter, but had a new calf to care for. We suggest that the greater impact on the behaviour and personality of surviving herd members following the deaths of calves, compared to an adult member, attests to the significance of the unifying role played by calves within an elephant herd.
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... The existence of salivary pheromones could explain trunk-to-mouth exchanges frequently observed between elephants [103,104]. Elephants also exhibit a variety of chemosensory behaviors when examining a dead conspecific, although we have little understanding of what information they might be gathering [86,[105][106][107]. The ubiquity of intraspecific chemical signaling by elephants challenges our imagination as to the types of information they are gathering. ...
... The sound of angrily buzzing bees evokes alarm calls and avoid behavior in elephants [105]. The smell of honey is associated with bees, and honey odor couples with bee buzzing sounds may indicate danger to elephants [106]. The multimodal signal may be more meaningful and thus resistant to habituation compared to single modality signals [107]. ...
Simple Summary
Among all taxa, messages transmitted via chemical signals are the oldest and most universal. For Asian and African elephants, odors convey information between individuals, and between elephants and their environment. Pheromones are chemical signals used within a species and while thousands of pheromones have been identified for insects, only a few dozen have been elucidated in mammals and other vertebrates. Amazingly, two pheromones are known for Asian elephants: one signals receptivity in females and the other a heightened reproductive state in males. The elephant trunk serves numerous functions including detecting airborne odors and transporting chemicals in substrates such as urine to be detected by multiple sensory systems. Obvious trunk behaviors provide clear means to assess the interest of elephants in scent sources. Thus, elephants can serve as a model system for investigating chemical signaling. Prior to the 21st century, research on elephant chemical signaling focused on within species communication. In the 21st century, these studies expanded with increasing fieldwork. Studies also revealed the use of odors to detect threats and forage for food. Chemical signaling in elephants remains a bouquet for further exploration with promising applications for the conservation of wild elephants and the management of elephants in human care.
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. 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.
... This point is echoed by Bercovitch, who notes that the comparative thanatology literature is exclusively composed of reports concerning social animals, and that there are no descriptions of reactions to the dead in solitary species, such as koalas or moose (Bercovitch 2020, p. 23). It has also been argued that the fission-fusion dynamics that characterise some social species could be crucial in the development of a CoD, insofar as these societies require the individuals to be constantly monitoring and updating the status of other individuals in the group (Goldenberg and Wittemyer 2020;Piel and Stewart 2015). ...
... Trapanese et al. (2020) report on a male capuchin monkey who inspected the anus of a stillborn infant and, immediately after, that of a live infant, possibly as a way of comparing olfactory cues. For elephants, smell may be the most salient characteristic of corpses, since they appear able to distinguish conspecific from heterospecific bones using olfaction and have also been witnessed smelling the ground where a corpse has decayed (Goldenberg and Wittemyer 2020). ...
Comparative thanatologists study the responses to the dead and the dying in nonhu-man animals. Despite the wide variety of thanatological behaviours that have been documented in several different species, comparative thanatologists assume that the concept of death (CoD) is very difficult to acquire and will be a rare cognitive feat once we move past the human species. In this paper, we argue that this assumption is based on two forms of anthropocentrism: (1) an intellectual anthropocentrism, which leads to an over-intellectualisation of the CoD, and (2) an emotional anthropocentrism, which yields an excessive focus on grief as a reaction to death. Contrary to what these two forms of anthropocentrism suggest, we argue that the CoD requires relatively little cognitive complexity and that it can emerge independently from mourning behaviour. Moreover, if we turn towards the natural world, we can see that the minimal cognitive requirements for a CoD are in fact met by many nonhuman species and there are multiple learning pathways and opportunities for animals in the wild to develop a CoD. This allows us to conclude that the CoD will be relatively easy to acquire and, so, we can expect it to be fairly common in nature.
... The references are in accordance with the ones suggested in the articles tools, except the references of 6 articles (Bastille-Rousseau et al. (2019) [5], [6], [90], Petracca et al. (2019) [2], Schiffmann et al. (2019) [106] and Watson et al. (2019) [110]), that are in accordance with the date of their first publication. Thus, the dates of the articles mentioned here are between July 2015 and December 2019. ...
... The [90] review highlights the behaviour of these individuals towards their dead. Touch and certain approaches to carcasses, regardless of the state of decomposition, as well as visiting carcasses and demonstrating extensive investigative behaviour, temporal gland streaming, selfdirected behaviour and intense social interactions among the living near carcasses prove the interest that elephants have in their deceased members and the strength of these social groups. ...
... For example, ants, as eusocial insects, may exhibit affective dimensions, such as pheromone-driven states (Jackson and Morgan, 1993;von Thienen et al., 2014), but likely lack reflective or narrative dimensions entirely. On the other hand, elephants may possess affective and intersubjective dimensions (Byrne et al., 2008;Hope et al., 2025) alongside something like narrative capacities, evidenced by their long-term memory and recognition of death (Bradshaw, 2009;Goldenberg and Wittemyer, 2020). Therefore, while some animals express a broad array of dimensions, others may operate effectively with a minimal set of aspects. ...
In the last decades, research on animal consciousness has advanced significantly, fueled by interdisciplinary contributions. However, a critical dimension of animal experience remains underexplored: the self. While traditionally linked to human studies, research focused on the self in animals has often been framed dichotomously, distinguishing low-level, bodily, and affective aspects from high-level, cognitive, and conceptual dimensions. Emerging evidence suggests a broader spectrum of self-related features across species, yet current theoretical approaches often reduce the self to a derivative aspect of consciousness or prioritize narrow high-level dimensions, such as self-recognition or metacognition. To address this gap, we propose an integrated framework grounded in the Pattern Theory of Self (PTS). PTS conceptualizes the self as a dynamic, multidimensional construct arising from a matrix of dimensions, ranging from bodily and affective to intersubjective and normative aspects. We propose adopting this multidimensional perspective for the study of the self in animals, by emphasizing the graded nature of the self within each dimension and the non-hierarchical organization across dimensions. In this sense, PTS may accommodate both inter- and intra-species variability, enabling researchers to investigate the self across diverse organisms without relying on anthropocentric biases. We propose that, by integrating this framework with insights from comparative psychology, neuroscience, and ethology, the application of PTS to animals can show how the self emerges in varying degrees and forms, shaped by ecological niches and adaptive demands.
... However, these data were retrospective and compared females when they did and did not have an infant; it is not possible to determine whether the change in behaviour was due to grief or an external environmental or other social variable [18]. Similarly, elephants (Loxodonta africana) who visit conspecific carcasses may show self-directed behaviours and temporal gland streaming, which may indicate an emotional response [19], but we are not aware of data showing that this occurs away from the carcass, which would indicate a grief response rather than a response to the stimulus of the carcass. Finally, there is some evidence that companion animals' behaviour shows grief-like changes after a co-residing conspecific has died, including less time playing, more time resting, and diminished appetite [20,21], but these survey data are influenced by the owners' perceptions and do not account for the change in the social environment of the bereaved individual. ...
Although it is understood that all humans grieve the death of close social partners, little empirical research has addressed animals’ responses to death. In this study, we collected quantitative data on the behaviour of 11 bereaved rhesus macaque (Macaca mulatta) mothers at Cayo Santiago to the natural deaths of their infants and matched, non-bereaved controls. Our research focused on behavioural signs of grief, including loss of appetite, lethargy, increased stress and social withdrawal, highlighting that such responses are documented in the human literature, but could be found in mammalian taxa. Using mixed models, we found that, contrary to prediction, bereaved mothers spent less time resting than the non-bereaved control females in the first two weeks after their infants’ deaths. There were no other behavioural markers of grief. We conclude that mothers showed a short-term behavioural response to their bereavement that does not match human’s prolonged ‘despair’ grief. We propose that mothers’ behavioural responses might be a form of ‘protest’ grief, as is seen in primate infants when separated from mothers and in humans, or do not grieve. We hope to advance the field of comparative thanatology by providing a framework and novel predictions for future studies in this area.
... Por lo tanto, no es sorprendente que muestren interés en los cadáveres, ya sean frescos o en un alto estado de descomposición, y que, a menudo, se observen comportamientos como el contacto físico, la exploración e incluso la agresión. Cabe señalar que algo aún más notable es que su fascinación puede persistir durante años (Goldenberg et al., 2020). ...
Este documento de investigación estudia la muerte, la enfermedad y el sufrimiento en el marco de las ciencias de la complejidad y en diálogo con las ciencias de la salud buscando establecer el sentido y cuidado de la vida.
... The participants were informed that she carried the body for two days, only setting it down to eat. After discussing the photos with the participants, they were told that elephants have been reported to approach dying companions and touch them, both before and after death, including helping their dying companions and guarding the dead bodies (Douglas-Hamilton et al. 2006;Goldenberg & Wittemyer 2020). Two videos were also shown in the cognition-emotion program, one where a mother elephant was trying to wake up her dead calf, and the other where elephants were helping a calf cross a river. ...
We evaluated whether the inclusion of elephant cognition in an education program resulted in changes in attitudes towards elephants and elephant conservation and the effect of rurality, gender, and age on attitudinal change. The study group was comprised of villagers living in and around Bannerghatta National Park. We found that adolescents had more favourable attitudes towards Asian elephants than adults, and participants in urban areas had more favourable attitudes than those in rural areas. No gender differences were observed. Overall, using elephant cognition as an educational tool did not result in more favourable attitudes towards elephants and elephant conservation.
... Additionally, their long-term memory skills help them to navigate through the landscape. Family herds are led by matriarchs who have a precise spatial memory allowing them to find water sources in times of drought (Polansky et al. 2015), but also the carcasses of lost relatives which elephants have been observed to revisit (Goldenberg and Wittemyer 2020). Elephants' long lifespan, their high intelligence, sociality and individuality makes it difficult to generalise elephant movements but also an enriching field of study. ...
Wildlife corridors are seen as essential environmental infrastructure guaranteeing species connectivity and biological diversity in contemporary conservation landscapes. Harking back to recent social science literature on infrastructure in general and environmental infrastructure in particular this contribution will analyze one contested wildlife corridor as part of a large conservationist project, the giant Kavango-Zambezi Transfrontier Conservation Area (KAZA TFCA), founded in 2011, and with 520,000 km2 the world’s largest conservation area. Using elephant corridors as an example, we will examine how they come into being and how they are governed. Only at first sight wildlife corridors appear as something entirely natural, constituted by the daily or seasonal movements of large herbivores. Indeed, elephants for example, produce paths through the savannah that are well visible in the landscape. But to make them wildlife corridors in conservation projects human action is needed: we will analyse wildlife corridors as hybrids of wildlife produced paths, administrative action and modern technologies of wildlife monitoring. As we will show in some detail focusing on one important Namibian wildlife corridor, the so-called Sobbe corridor, the human-elephant conflict is a real challenge leading to local resistance to corridors. Surprisingly, though, conflict is more frequent along the linear settlements close to roads, than on wildlife corridors, instances of human-elephant conflict is a prominent topic linked to corridors. The most significant challenge to corridors though comes from the rapid expansion of agricultural fields as well as from local conflicts over land ownership.
... Another behavior that may be suggestive of a long-term social memory capacity in elephants is the grief-like behaviors that elephants have been observed engaging in following the death of conspecifics. For example, in the wild, all three species of elephants have been observed investigating, touching, and sniffing carcasses, and revisiting them repeatedly over the course of a few minutes to a few months (Douglas-Hamilton et al., 2006;Goldenberg & Wittemyer, 2020;Hawley et al., 2018;McComb et al., 2006;Merte et al., 2009;Payne, 2003;Pokharel et al., 2022;Sharma et al., 2020;Stephan et al., 2020). Captive Asian elephants also show changes in social and affiliative behaviors after the loss of a herd member (Rutherford & Murray, 2021). ...
Despite popular culture’s promotion of the elephant’s ability to “never forget,” there is remarkably limited empirical research on the memory capacities of any living elephant species (Asian, Elephas maximus; African savanna, Loxodonta africana; African forest, Loxodonta cyclotis). A growing body of literature on elephant cognition and behavioral ecology has provided insight into the elephant’s ability to behave flexibly in changing physical and social environments, but little direct evidence of how memory might relate to this flexibility exists. In this paper, we review and discuss the potential relationships between what we know about elephant cognition and behavior and the elephants’ memory for the world around them as they navigate their physical, social, and spatial environments. We also discuss future directions for investigating elephant memory and implications for such research on elephant conservation and human–elephant conflict mitigation.
... Por lo tanto, no es sorprendente que muestren interés en los cadáveres, ya sean frescos o en un alto estado de descomposición, y que, a menudo, se observen comportamientos como el contacto físico, la exploración e incluso la agresión. Cabe señalar que algo aún más notable es que su fascinación puede persistir durante años (Goldenberg et al., 2020). ...
En repetidas ocasiones, los filósofos han dejado claro que la fragilidad humana es evidente. La pandemia del COVID-19 ha puesto en evidencia lo expuesto anteriormente y lo ha hecho a nivel global. Aunque algunos gobernantes han declarado que la pandemia ha terminado, la verdad es que todavía está presente. La pandemia ha demostrado que, a menudo, los gobernantes priorizan la economía y ponen en riesgo la salud de las personas. Actualmente, estamos siendo testigos del fenómeno del COVID-19 de largo aliento, el cual todavía no ha sido comprendido en su totalidad por los tomadores de decisiones y los profesionales de la salud.
... Elephants hold significant intrinsic, biological, ecological and human cultural value. They possess unique genetic and physiological features [1], exhibit high levels of individual cognitive and emotional intelligence [2][3][4], maintain complex social behaviours and structures [5][6][7][8][9], serve as keystone species in their respective ecosystems [10][11][12], are highly valued by their local communities [13] and serve as flagship species for animal conservation [14]. ...
Artificial intelligence (AI) and machine learning (ML) present revolutionary opportunities to enhance our understanding of animal behaviour and conservation strategies. Using elephants, a crucial species in Africa and Asia’s protected areas, as our focal point, we delve into the role of AI and ML in their conservation. Given the increasing amounts of data gathered from a variety of sensors like cameras, microphones, geophones, drones and satellites, the challenge lies in managing and interpreting this vast data. New AI and ML techniques offer solutions to streamline this process, helping us extract vital information that might otherwise be overlooked. This paper focuses on the different AI-driven monitoring methods and their potential for improving elephant conservation. Collaborative efforts between AI experts and ecological researchers are essential in leveraging these innovative technologies for enhanced wildlife conservation, setting a precedent for numerous other species.
... Por lo tanto, no es sorprendente que muestren interés en los cadáveres, ya sean frescos o en un alto estado de descomposición, y que, a menudo, se observen comportamientos como el contacto físico, la exploración e incluso la agresión. Cabe señalar que algo aún más notable es que su fascinación puede persistir durante años (Goldenberg et al., 2020). ...
Este Working Paper estudia los desastres y calamidades a la luz de la vida, no solamente de la tragedia. Los sismeas vivos no solamente son resilientes; mucho mejor, tienen la capacidad de reprarse a sí mismo, y explorar incesantemente posiblidades.
... Stillborn babies and dead infants often elicit maternal care that can persist for days, weeks, or even months in non-human primates (see Anderson 2011;Watson and Matsuzawa 2018;Gonçalves and Carvalho 2019), but similar responses have also been observed in other mammals such as elephants (Goldenberg and Wittemyer 2020), and cetaceans and other aquatic mammals (Reggente et al. 2016). These behaviors, which can be associated with the hormonal state of the mother and/or her emotional attachment to the infant (Biro et al. 2010), possibly reflect a lack of understanding by the mother about the death of the baby (Hrdy 1979). ...
Comparative thanatology includes the scientific study of death in non-human animals, which involves emotional, social, and exploratory responses of individuals and groups towards corpses. Stillborn babies and dead infants often elicit maternal and alloparental care that can persist for days, weeks, or even months, especially in primates. After this period, cannibalistic acts can occur not only by groupmates but also by the mother. Such cannibalism has been reported both in captive and wild primate groups, suggesting that the phenomenon is evolutionary adaptive. Here, we report a case in drills (Mandrillus leu-cophaeus), a largely unstudied monkey species. We collected data from the birth to the death of the infant on maternal and alloparental care towards the newborn across three phases: pre-mortem, post-mortem, and post-mortem cannibalism. The mother maintained high levels of grooming after the infant's death. Both the mother and other group members interacted with the dead baby by trying to engage its gaze. Two days after the death, the mother started to eat the corpse until it was almost completely consumed; there was no sharing with other group members. Although we cannot draw firm conclusions about potential benefits of the mother's behavior, this observation on drills adds a piece to the puzzle of thanatological behaviors and cannibalism in primates.
... Indeed, there is preliminary evidence in support of elephants as a relevant animal model for cultural evolution. For instance, elephants exhibit behaviors that are indicative of self-awareness and potentially the existence of ''theory-of-mind,'' including the attribution of mental or biological states to others (174), displays of empathy (96,98), mirror self-recognition (267,268), and sensitivity to the remains of deceased conspecifics (95,269). In addition, elephants display tool use, which is another significant marker of cultural evolution (270,271). ...
Humans are unique in their sophisticated culture and societal structures, their complex languages, and their extensive tool use. According to the human self-domestication hypothesis, this unique set of traits may be the result of an evolutionary process of self-induced domestication, in which humans evolved to be less aggressive and more cooperative. However, the only other species that has been argued to be self-domesticated besides humans so far is bonobos, resulting in a narrow scope for investigating this theory limited to the primate order. Here, we propose an animal model for studying self-domestication: the elephant. First, we support our hypothesis with an extensive cross-species comparison, which suggests that elephants indeed exhibit many of the features associated with self-domestication (e.g., reduced aggression, increased prosociality, extended juvenile period, increased playfulness, socially regulated cortisol levels, and complex vocal behavior). Next, we present genetic evidence to reinforce our proposal, showing that genes positively selected in elephants are enriched in pathways associated with domestication traits and include several candidate genes previously associated with domestication. We also discuss several explanations for what may have triggered a self-domestication process in the elephant lineage. Our findings support the idea that elephants, like humans and bonobos, may be self-domesticated. Since the most recent common ancestor of humans and elephants is likely the most recent common ancestor of all placental mammals, our findings have important implications for convergent evolution beyond the primate taxa, and constitute an important advance toward understanding how and why self-domestication shaped humans' unique cultural niche.
... Elephants also exhibit displacement activity and auto-related behaviours, but this index has not been fully explored. Several authors discussed trunk manipulation/ self-touching behaviours in free-ranging African elephants (Loxodonta africana), and attributed this to states of internal conflict (Douglas-Hamilton, 1975) or situations of apprehension and/or distress (Mason and Veasey, 2010;Douglas-Hamilton et al., 2006;Goldenberg and Wittemyer, 2020). However, the connection between SDBs and potential implications of the elephant's mental state in terms of welfare has not been made to date. ...
Captive African elephants used in the tourism industry face numerous welfare issues which are often stress related and linked to high numbers of tourists or human-elephant interactions. Elephant welfare is commonly assessed by quantifying faecal glucocorticoid metabolite (fGCM) concentrations, and monitoring stereotypic behaviour, which are either costly or problematic in identifying underlying causes. Self-directed behaviours (SDBs), a form of displacement activity, have been substantially linked with stress and anxiety in primates, and could be a potential but not yet utilised behavioural marker in elephants. We thus explored the usage of several trunk, tail, and body related SDBs in a group of 7 semi-captive African elephants maintained at the Knysna Elephant Park, a tourist destination offering a variety of close contact experiences. Using continuous behavioural sampling (focal following), SDBs were recorded as they occurred, along with the numbers of nearby tourists (<5 m to focal; with 3 levels: None, Low 1-5, and High >5), and the nature of the tourist interaction (5 levels: None, Feed, Touch, Walk, and Ride). We then compared SDB rates with fGCM concentrations (n = 115) determined following observations (1 day and 2 days after behavioural sampling). Data were analysed using Generalized Linear Mixed effects Models. SDB rates (p/min) significantly increased during elephant Walks (p < 0.001) and Rides (p < 0.001), indicating a correlation with their usage during potentially stressful scenarios. The Touch interaction significantly decreased SDBs (p < 0.05), whilst the Feed interaction had a non-significant effect (p > 0.05), which could indicate the element of control in terms of perceived stress is an important component in welfare, as elephants were free to move away during these interactions. Interestingly, SDBs significantly decreased when tourist numbers were High (p = < 0.05), potentially due to greater quantities of high value food. Additionally, no correlation was found between SDBs and fGCM concentrations determined on day one (p > 0.05), and day two (p > 0.05). Hence, SDBs may be correlated more with anxiety or lower-level stress, which is not significant enough to activate GC production, and may therefore act as a coping strategy utilised to maintain physiological homeostasis during anxiety-inducing situations. Additional research would benefit from coupling SDB observations with different forms of physiological assessments to better understand the internal motivations, and formally establish SDBs as a reliable, cost-effective, and non-invasive welfare index to identify stressed individuals in real time and ensure optimum husbandry.
... Também pombos apresentam distress em resposta ao distress percebido num conspecífico (Waal, 2008) e cães conseguem sentir empatia pelos donos quando choram e, nessa situação, tentam acudi-los rapidamente (Sanford et al., 2018). Os elefantes tranquilizam companheiros angustiados e são conhecidos por apoiar outros mais fracos para ficar de pé (Waal, 2008) e apresentar sinais de luto na presença de um familiar morto (Goldenberg & Wittemyer, 2020). Li et al. (2018) observaram que existe contágio social vicário de dor entre ratos familiares, dependente de expressões de dor, como também do tempo que as díades gastam em comunicações sociais. ...
O presente estudo consiste numa abordagem teórica acerca das interacções entre humanos e animais não-humanos, em que se utiliza o contributo da antrozoologia e do bem-estar animal para compreender os factores que as moldam, e se expõe uma compilação de alguns estudos para exemplificar formas de as melhorar no contexto pecuário. Na segunda-parte sintetizam-se diferentes abordagens de ética animal que podem contribuir para a compreensão da natureza de algumas interacções e analisam-se três casos de estudo. Esta análise crítica, bem como a discussão dos elementos fundamentais que moldam as interacções, permite a sistematização de possíveis soluções para melhorar o papel do tratador/operador na convivência com os animais.
Conclui-se que uma má qualidade na interacção entre humanos e animais dificulta o maneio com os animais e contribui para o empobrecimento da qualidade de vida de ambos. Este trabalho sublinha a importância que o perfil pessoal, as competências e a formação técnica e pessoal do tratador têm para cumprir as suas tarefas profissionais com critérios elevados de bem-estar animal, assim estabelecendo uma interacção com mútuos benefícios.
Palavras-chave: interacções entre humanos e animais (HA); antrozoologia, bem-estar animal, ética animal, stockmanship
... Elephants appear to have learned to discern between human ethnic groups that vary in their level of threat toward elephants McComb et al., 2014). Elephants also seem to be unique among non-human animals in that they may exhibit behaviors related to "theory of mind", demonstrating selfawareness (Plotnik et al., 2010), cooperation with one another (Plotnik et al., 2011), mourning-like behavior (Goldenberg and Wittemyer, 2020), empathy , and consolation (Plotnik and de Waal, 2014). ...
Elephants are large-brained, social mammals with a long lifespan. Studies of elephants can provide insight into the aging process, which may be relevant to understanding diseases that affect elderly humans because of their shared characteristics that have arisen through independent evolution. Elephants become sexually mature at 12 to 14 years of age and are known to live into, and past, their 7 th decade of life. Because of their relatively long lifespans, elephants may have evolved mechanisms to counter age-associated morbidities, such as cancer and cognitive decline. Elephants rely heavily on their memory, and engage in multiple levels of competitive and collaborative relationships because they live in a fission-fusion system. Female matrilineal relatives and dependent offspring form tight family units led by an older-aged matriarch, who serves as the primary repository for social and ecological knowledge in the herd. Similar to humans, elephants demonstrate a dependence on social bonds, memory, and cognition to navigate their environment, behaviors that might be associated with specializations of brain anatomy. Compared with other mammals, the elephant hippocampus is proportionally smaller, whereas the temporal lobe is disproportionately large and expands laterally. The elephant cerebellum is also relatively enlarged, and the cerebral cortex is highly convoluted with numerous gyral folds, more than in humans. Last, an interesting characteristic unique to elephants is the presence of at least 20 copies of the TP53 tumor suppressor gene . Humans have only a single copy. TP53 encodes for the p53 protein, which is known to orchestrate cellular response to DNA damage. The effects of these multiple copies of TP53 are still being investigated, but it may be to protect elephants against multiple age-related diseases. For these reasons, among others, studies of elephants would be highly informative for aging research. Elephants present an underappreciated opportunity to explore further common principles of aging in a large-brained mammal with extended longevity. Such research can contribute to contextualizing our knowledge of age-associated morbidities in humans.
... Tursiops truncatus, Stenella coeruleoalba, Sousa chinensis) often stay close to dead conspecifics, and they may carry and keep them at the water surface, presumably to help them breathe (Bearzi et al. 2018;Reggente et al. 2018). The most commonly recorded behaviours in elephants (Loxodonta africana, Elephas maximus) appear to be investigative, but elephants may also push and pull dying or recently deceased individuals, attempting to make them stand up; they sometimes vocalise or mount a corpse, and in rare cases they have been seen to cover it with vegetation (Moss 1988;Payne 2003;Douglas-Hamilton et al. 2006;Bercovitch 2020;Goldenberg & Wittemyer 2020;Sharma et al. 2020). Data in other vertebrates remain scarce and physical contact with the dead appear to be more limited. ...
It was long assumed that only humans can distinguish the living from the dead. Renewed interest in this question over the last decade has led several authors to assert that non-human primates are also aware of death. We investigate this issue by comparing the behaviours of monkeys and great apes toward helpless conspecifics, basing our analysis on published reports. We first examine the behaviours of mothers towards the body of their dead offspring. They may carry the corpse for days or more before abandoning it. They groom, inspect and protect it, sometimes allowing group members to explore it, and rare cases of cannibalism have been reported. No significant difference is observed in the way that monkeys and great apes treat the bodies of infants. We then examine responses to collapsed (still able to move and react) and inanimate (unresponsive or dead) conspecifics. Monkeys and great apes guard, care for and inspect their helpless partners, and also manipulate and mobilise them. Through these actions, individuals may inform themselves about the state of their partners, test their responsiveness and/or attempt to rouse them. It appears that only chimpanzees and gorillas show violent action such as display behaviours and the rough treatment of bodies. They can also make distress calls, and periods of “stunned silence” sometimes occur in chimpanzees, indicating that they are experiencing intense emotion. Finally, we argue that while both monkeys and great apes detect body dysfunction through the victims’ inability to wake up and move, only great apes can understand that something serious has happened. The signs of emotional disturbance reported in them indicate that they may believe that inanimate conspecifics have entered a state of “dormancy”, meaning that they are unlikely to regain wakefulness. However, there is no evidence that any non-human primates are aware of mortality.
... . Species facing the necessity to manoeuvre short-term social relationships in fission-fusion systems seem to show particular interest in dead conspecifics (chimpanzees, Pan troglodytes: Stewart et al., 2012;savannah elephants, Loxodonta africana africana: Douglas-Hamilton et al., 2006;cetaceans: Bearzi et al., 2018), most likely to update social information (Anderson, 2016). For instance, savannah elephants, who depend on social information transfer and who possess many differentiated social relationships throughout life (McComb et al., 2001;Foley, 2002), have been repeatedly reported to approach, investigate and to touch deceased conspecifics, even when they have not previously been closely bonded to them (Goldenberg & Wittemyer, 2019). ...
The increased attention towards deceased conspecifics in various social animal species is one of the most intriguing conundrums in animal behaviour. The factors that might explain the observed behavioural variation amongst individuals remain nebulous. Here we analyse forest elephants’ ( Loxodonta africana cyclotis ) responses to a poached adult male conspecific, using remote camera trapping during a period of eight months. After completely avoiding the carcass site for over a week, females and males substantially differed in behavioural responses. Males consistently stayed longer around the remains, showed signs of increased arousal, interacted with the dead body, and twisted trunks with each other. Females, in contrast, were more passively explorative and preferred to visit the site without their dependent offspring. Findings show a previously unknown sexual-dimorphism in forest elephant behaviour towards a poached conspecific and raise the possibility that individuals might be able to infer further context-specific information about the event.
African elephants (Loxodonta africana) are the largest extant terrestrial mammals, with bodies containing enormous quantities of nutrients. Yet, we know little about how these nutrients move through the ecosystem after an elephant dies. Here, we investigated the initial effects (1–26 months postmortem) of elephant megacarcasses on savanna soil and plant nutrient pools in the Kruger National Park, South Africa. We hypothesized that (H1) elephant megacarcass decomposition would release nutrients into soil, resulting in higher concentrations of soil nitrogen (N), phosphorus (P), and micronutrients near the center of carcass sites; (H2) carbon (C) inputs into the soil would stimulate microbial activity, resulting in increased soil respiration potential near the center of carcass sites; and (H3) carcass-derived nutrients would be absorbed by plants, resulting in higher foliar nutrient concentrations near the center of carcass sites. To test our hypotheses, we identified 10 elephant carcass sites split evenly between nutrient-poor granitic and nutrient-rich basaltic soils. At each site, we ran transects in the four cardinal directions from the center of the carcass site, collecting soil and grass (Urochloa trichopus, formerly U. mosambicensis) samples at 0, 2.5, 5, 10, and 15 m. We then analyzed samples for C, N, P, and micronutrient concentrations and quantified soil microbial respiration potential. We found that concentrations of soil nitrate, ammonium, δ15N, phosphate, and sodium were elevated closer to the center of carcass sites (H1). Microbial respiration potentials were positively correlated with soil organic C, and both respiration and organic C decreased with distance from the carcass (H2). Finally, we found evidence that plants were readily absorbing carcass-derived nutrients from the soil, with foliar %N, δ15N, iron, potassium, magnesium, and sodium significantly elevated closer to the center of carcass sites (H3). Together, these results indicate that elephant megacarcasses release ecologically consequential pulses of nutrients into the soil which stimulate soil microbial activity and are absorbed by plants into the above-ground nutrient pools. These localized nutrient pulses may drive spatiotemporal heterogeneity in plant diversity, herbivore behavior, and ecosystem processes.
Whereas humans exhibit emergency responses to assist unconscious individuals, how nonhuman animals react to unresponsive conspecifics is less well understood. We report that mice exhibit stereotypic behaviors toward unconscious or dead social partners, which escalate from sniffing and grooming to more forceful actions such as mouth or tongue biting and tongue pulling. The latter intense actions, more prominent in familiar pairs, begin after prolonged immobility and unresponsiveness and cease when the partner regains activity. Their consequences, including improved airway opening and clearance and accelerated recovery from unconsciousness, suggest rescue-like efforts. Oxytocin neurons in the hypothalamic paraventricular nucleus respond differentially to the presence of unconscious versus active partners, and their activation, along with oxytocin signaling, is required for the reviving-like actions. This tendency to assist unresponsive members may enhance group cohesion and survival of social species.
The mass influx of Rohingyas to Cox’s Bazaar refugee camps in 2017 blocked the only migratory corridor used by the critically endangered Asian Elephants to travel between their habitats in Bangladesh and Myanmar. In an unforeseen collision between Rohingya refugees and Asian elephants, humanitarian and conservation actors have found themselves at an impasse—protecting one group threatens the existence of another. Temporary interventions were executed to mitigate the conflict and save human lives, but without extensive, long-term solutions, the elephants risk losing genetic diversity and even extinction. Analyzed with posthumanist ethics, on the ground realities of inadequate conservation measures and a deficient legislative landscape, it becomes apparent that a new stakeholder framework must be proposed, inclusive of all populations that are directly or indirectly impacted, tracing the conflict back to a central actant.
This case study, inspired by the work of Kaswan and Roy, provides critical insights into the thanatological behaviour of elephants, specifically in the North Bengal landscape. It serves as an important resource for private sanctuary managers, elephant handlers, and conservationists worldwide, raising awareness of elephants’ intelligence, emotional depth, and strong bonds with their herd members. The study reveals a unique behaviour among elephants in a specific area, where they bury deceased calves in 2-foot-deep irrigation drains within tea gardens. This region, featuring fragmented forests, tea estates, and small towns, serves as a natural corridor for elephants. The calves are buried in a ‘leg-upright’ position, indicative of how the herd uses their trunks and legs to manipulate the bodies. Two calves died from fungal infections, while three faced respiratory failure, yet all were buried similarly. Contusion marks suggest the herd dragged the calves over a distance before burial, highlighting their collective handling of the bodies. This burial behaviour is likely driven by several factors, including the particularly strong mother-calf bond seen in elephants which may motivate the herd to engage in mourning and burial. The unique environmental conditions in North Bengal, combined with the elephants’ complex social and emotional behaviours, create a remarkable example of interspecies empathy and care, offering valuable lessons for global conservation and elephant management efforts.
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This short paper is intended to alert our colleagues to the existence of The Elephant Ethogram: A Library of African Elephant Behaviour. It describes its purpose, form and scope, and appeals for contributions of undocumented, rare, novel or cultural Loxodonta africana behaviour. We do not present descriptions of behaviours, methodologies, results or discussion; these may be found online within The Elephant Ethogram. The Elephant Ethogram is an ElephantVoices initiative to document the complex, diverse and nuanced repertoire of behaviour and communication of African savannah elephants (L. africana). In a unique, user-friendly, fully searchable and publicly accessible database, The Elephant Ethogram chronicles the rich postural, gestural, tactile, chemical and acoustic communication and behaviour of Africa's savannah elephants. It includes commonly displayed, unusual, novel and culturally learned behaviours, as well as those expressed in response to people. The Elephant Ethogram is based on published descriptions of behaviour and the decades of behavioural studies and photographic, acoustic and video-graphic collections from Amboseli National Park (NP), the Maasai Mara ecosystem (Mara), Kenya, and Gorongosa National Park, Mozambique conducted by ElephantVoices. It is also built from behaviour captured for documentaries filmed in the Mara and Gorongosa and video clips of unusual behaviour collected by colleagues and members of the general public. The Elephant Ethogram replaces ElephantVoices' online Elephant Gestures and Elephant Calls Databases originally developed in 2003 and revamped in 2011 (Poole 2011; Poole and Granli 2011), that were based on our elephant studies in Amboseli National Park, Kenya between 1975 and 2009 (Poole 1987; Poole et al. 1988; Poole 1989a; Poole 1989b; Poole et al. 2005) and the work of other scientists (Douglas-Hamilton 1972; Berg 1983; Moss 1983; Kahl and Armstrong 2000; Kahl and Armstrong 2002). Between 2011 and 2019 we carried out elephant behaviour studies and conservation projects in the Mara, and Gorongosa NP, during which we completed elephant, field notes, images and videos of the behaviour of known individuals. Furthermore, in a ground-breaking collaboration with copyright owners Off the Fence, Gorongosa Media Project and Bob Poole Films, hundreds of hours of raw footage of elephants, originally shot for documentaries in Gorongosa and the Mara, were granted to ElephantVoices for use in science and education. Since we collaborated with the filmmakers on site, the footage primarily depicts known individuals. In 2020 we collected additional footage of behaviour in Amboseli National Park. The Elephant Ethogram combines and significantly improves the structure and functionality of the original databases, includes hundreds of additional behaviours, 2,400 annotated video clip examples from three populations, higher-resolution images, additional audio files. Elephant behaviour has been documented by hunters, naturalists and scientists for hundreds of years starting with the earliest scholarly notes of Aristotle (1862 translation) to those of Darwin (1872) to Kühme’s (1962; 1963) research on captive African elephants. The study of free-ranging elephants by Douglas-Hamilton (1972) stimulated the work of many who followed, including the five decades-long work of Moss and her colleagues, and our own. From hundreds of published studies, we know that savannah elephants show great richness, variation and flexibility in their behaviour. Some of these publications have described elephant behaviours relevant to the aims of their particular study (Moss 1983 (oestrus); Poole 1987 (musth); O’Connell-Rodwell et al. 2011 (male-male relationships); Goldenberg and Wittemyer 2020 (death)), but only Kahl and Armstrong’s work in Hwange National Park, Zimbabwe (2000; 2002) and our own in Amboseli National Park (Poole 2011; Poole and Granli 2011) aimed to document the full repertoire of behaviour of the species. We worked closely with Kahl to share data and to agree on terminology and definitions. Until his untimely death in 2012 Kahl’s plan was to publish a detailed elephant ethogram. The construction of “exclusive ethograms” to describe a species' behaviour or activity patterns is commonly used in behavioural studies, where the ethogram focuses on the behaviours of interest. It is more unusual to find catalogues that attempt to produce an “exhaustive” ethogram of all known behaviours of a given species. One example is the work of Nishida et al. (1999) on chimpanzees, Pan troglodytes, which aims to provide an exhaustive list of behaviours. This body of work defines 515 behaviours, recording whether they were idiosyncratic, limited to a small group, to one population or were, to a greater or lesser extent, cross-cultural. Another example is the work of Bolgan et al. (2014) on the Arctic charr, Salvelinus alpinus, a fish species. A draw-back of these studies is that they are non-searchable, written documents without video-graphic documentation. Nishida et al. (2010) solved this problem for their study of chimpanzees by publishing a book with an accompanying CD. Very few studies; however, have attempted to produce an exhaustive, searchable web-based ethogram of a species with video examples of behaviour. One example is Mouse Ethogram: An Ethogram for the Laboratory Mouse developed in the Stanford Medical School in the Laboratory of Joseph Garner. Since this study was carried out in a captive environment it is unlikely to be exhaustive for the species. African elephants (Savannah and Forest) are among the most socially complex non-human species (Moss and Poole 1983; McComb et al. 2000; Archie et al. 2005; Wittemyer et al. 2005), as well as one of the more heavily exploited (Meredith 2001; Wittemyer et al. 2014). As scientists continue to document their extraordinary behaviour, elephants are increasingly impacted by humans to the point where their behaviour is notably affected (Douglas-Hamilton et al. 2005; Gaynor et al. 2018; Wall et al. 2021) and their future survival endangered (Wittemyer et al. 2014; Hart et al. 2021).
We note thanatological behavioural displays by clan members of a deceased spotted hyaena, Crocuta crocuta (Erxleben, 1777) in a fenced reserve. A matriarch had a lethal altercation with lions, Panthera leo (Linnæus 1758), on 10.V.2022. The carcass was monitored with a camera trap that recorded videos until the carcass was completely decayed. Four instances of clan members were recorded observing and sniffing the genitalia of the deceased hyaena over a period of six days. Within one month, the clan killed the deceased’s cubs and moved their den location. We recommend further research be conducted into thanatological behaviours of spotted hyaenas as these behaviours could affect ethical considerations of management practices.
Rampant environmental changes and forest destruction push elephants, both Asian and African, to explore human spaces to fulfil their dietary and ecological requirements and, consequently in shared spaces many ‘novel’ elephant behaviors come into the limelight. Elephant calf burial is reported in African literature but remains absent from the Asian context. We report calf burials by Asian Elephants in the eastern Himalayan floodplains of the northern Bengal landscape. The study area consists of fragmented forests, tea estates, agricultural lands, and military establishments. Tea estates form the majority of elephant corridors, and we explain the burial strategy of elephants in the irrigation drains of tea estates. We present five case reports of calf burials by elephants. We aimed to understand the perimortem strategy and postmortem behavior of the Asian Elephants. The major findings reflect that the carcasses were carried by trunks and legs for a distance before being buried in a ‘legs-upright-position’. We further investigated the underlying reason for calf deaths through postmortem examinations. Direct human intervention was not recorded in any of the five deaths. Through opportunistic observation, digital photography, fieldnotes, and postmortem examination reports, we suggest that the carcasses were buried in an abnormal recumbent style irrespective of the reason for the calf’s death. Through long-term observation, we further report that the elephants in this region clearly avoid the paths where carcasses were buried. We discuss and connect the literature of two distinct elephant species and also compare thanatological studies of other sentient nonhuman species.
African (Bantu) philosophy conceptualizes morality through ubuntu, which emphasizes the role of community in producing moral agents. This community is characterized by practices that respond to and value interdependence, such as care, cooperation, and respect for elders and ancestral knowledge. While there have been attributions of morality to nonhuman animals in the interdisciplinary animal morality debate, this debate has focused on Western concepts. We argue that the ubuntu conception of morality as a communal practice applies to some nonhuman animals. African elephant communities are highly cooperative and structured around elders; they alloparent, protect their communities, mourn their dead, and pass on cultural knowledge between generations. Identifying these as important moral practices, ubuntu provides a theoretical framework to expand our ethical concern for elephants to their communities. In practice, this will deepen our understanding of the wrongness of atrocities like culling for population management or trophy hunting.
Slowly reproducing and long-lived terrestrial mammals are often more at risk from challenges that influence fitness and survival. It is, therefore, important to understand how animals cope with such challenges and how coping mechanisms translate over generations and affect phenotypic plasticity. Rapidly escalating anthropogenic challenges may further diminish an animal’s ability to reinstate homeostasis. Research to advance insights on elephant stress physiology has predominantly focused on relative or comparative analyses of a major stress response marker, glucocorticoids (GCs), across different ecological, anthropogenic, and reproductive contexts. This paper presents an extensive review of published findings on Asian and African elephants from 1980 to 2023 (May) and reveals that stress responses, as measured by alterations in GCs in different sample matrices, often are highly dynamic and vary within and across individuals exposed to similar stimuli, and not always in a predictable fashion. Such dynamicity in physiological reactivity may be mediated by individual differences in personality traits or coping styles, ecological conditions, and technical factors that often are not considered in study designs. We describe probable causations under the ‘Physiological Dynamicity Model’, which considers context–experience–individuality effects. Highly variable adrenal responses may affect physiological plasticity with potential fitness and survival consequences. This review also addresses the significance of cautious interpretations of GCs data in the context of normal adaptive stress versus distress. We emphasize the need for long-term assessments of GCs that incorporate multiple markers of ‘stress’ and ‘well-being’ to decipher the probable fitness consequences of highly dynamic physiological adrenal responses in elephants. Ultimately, we propose that assessing GC responses to current and future challenges is one of the most valuable and informative conservation tools we have for guiding conservation strategies.
Many nonhuman animals have the emotional capacities to form caring relationships that matter to them, and for their immediate welfare. Drawing from care ethics, we argue that these relationships also matter as objectively valuable states of affairs. They are part of what is good in this world. However, the value of care is precarious in human-animal interactions. Be it in farming, research, wildlife ‘management’, zoos, or pet-keeping, the prevention, disruption, manipulation, and instrumentalization of care in animals by humans is ubiquitous. We criticize a narrow conception of welfare that, in practice, tends to overlook non-experiential forms of harm that occur when we interfere with caring animals. Additionally, we point out wrongs against caring animals that are not just unaccounted for but denied by even an expansive welfare perspective: The instrumentalization of care and caring animals in systems of use can occur as a harmless wrong that an approach purely focused on welfare may, in fact, condone. We should therefore adopt an ethical perspective that goes beyond welfare in our dealings with caring animals.
The elephants are the most important animal characters in Nirmal Ghosh’s novella River Storm, and they both metaphorically and literally permeate the entire story. This article explores how viewing these animals through the lens of sensory ecology enables more accurate identification of the ways in which the elephants’ olfactory, tactile, and auditory senses help in communicating with their environment and a better comprehension of the nonhuman perspective. It also tries to illustrate why understanding an elephant’s sensory capabilities is necessary to acknowledge the non-human perspective in a society where humans predominate. Drawing on the premises of Jakob Von Uexkull’s umwelt theory, this study acknowledges the elephants’ agency by contradicting humans’ generic conception of animals as objects or symbols or allegorical subjects in the epistemology of literary discourse about animals. To emphasize the elephant’s distinctive perception of the environment, the narrative specifically moves beyond the boundaries of human perception. Furthermore, it clarifies how an elephant interacts with their surroundings using their sensory faculties, how misinterpretations of their umwelt can result in a number of conflicts between humans and elephants, as well as how a clear understanding of the elephant world can result in a cordial relationship between humans and elephants.
Reporting reactions to death in nonhuman primates can provide valuable information for understanding the evolutionary origin of human ways of dealing with death. Although many studies have reported nonhuman primates' reactions to infant corpses, less is known about their reactions toward dead adults or adolescents. The deaths of adult primates, who usually form complex social relationships, may have social impacts. Here, we described in detail the reactions of Japanese macaques (Macaca fuscata) to the corpse of an adult female in a free-ranging multi-male multi-female group. We analyzed quantitative data on the whole group to determine how death-related behaviors were related to social relationships. Most group members in this case, including social partners of the deceased, exhibited no notable interest in the corpse. Only one adult female, who was not a grooming partner of the female before she died, touched and groomed the corpse. We examined four possible reasons why this female groomed the corpse: unawareness of death, learning about death, desire to consume insects, and reputation-building with other group members. This study highlights the potential value of closer examination of associations between reactions to dead adults or adolescents and social relationships before and after death in primate groups.
Primate thanatology, or the study of primate responses to dying and death, has become increasingly relevant in recent years. However, the number of reports remains small and the quality of published records is highly variable. Here, we extend the literature on comparative thanatology with observations on a population of the northern muriqui (Brachyteles hypoxanthus). We compiled all seven cases of muriqui responses to dead, dying, or abandoned infants observed by trained researchers of the Muriqui Project of Caratinga between 1998 and 2020. Four different adult females were observed carrying their dead offspring (n = 4) and one of these females was also observed carrying her dead grand-offspring (n = 1). Five of the seven cases involved dead newborns, one involved a newborn abandoned by its mother on the forest floor, and one involved a 1.6-year-old infant that was visibly ill and died after its mother left it on the ground. Dead newborns were carried for 1–3 days, and all cases occurred during the dry season months. No other age-sex classes were observed to interact with the dead or dying immatures. Our observations are consistent with hypotheses concerning the predominance of dead-infant carrying in other primates, and with the effects of climate on rates of corpse disintegration. They also show the value of long-term studies for obtaining and understanding anecdotal records of rare behavior.
We discussed the relation between human and non-human animals, a connection that is illustrated by Charles Darwin’s evolution theory, underpinned by modern genome sequencing techniques. Although differences between human and non-human animals may appear as more striking, similarities are much greater and more profound. These similarities go beyond anatomical and physiological features; they also include aspects as feelings and compassion with others. We then asked how Homo sapiens, despite being so similar, managed to sit on the top of the monkey rock, from where it dominates and exploits so many other species. The ability to form large groups of collaborating individuals, strongly guided by common believes and emotions, makes people so dominant over other animals. We took a closer look into the history of people, in particular veterinarians, caring for animals. The veterinary profession, as liberal profession, constantly tries to balance various interests. Every veterinarians is committed to respect ethical principles. Further research, education and specialization in animal welfare lead to a better understanding of how the welfare of animals shall be best assured. Caring for the health and welfare of animals and people is in the veterinary genes. A next step in the profession’s evolution could be a stronger competence to guide societal changes. Inspiring animal owners and caretakers to considering animals as, in the words of Charles Darwin, our fellow brethren and companions would be a great aspiration. No better recognition for Darwin, and tremendously helpful for all animal species, not the least our own.KeywordsEvolutionAnimal welfareEmpathyVeterinarianLiberal profession
Observations of some mammals and birds touching their dead provoke questions about the motivation and adaptive value of this potentially risky behaviour. Here, we use controlled experiments to determine if tactile interactions are characteristic of wild American crow responses to dead crows, and what the prevalence and nature of tactile interactions suggests about their motivations. In Experiment 1, we test if food or information acquisition motivates contact by presenting crows with taxidermy-prepared dead crows, and two species crows are known to scavenge: dead pigeons and dead squirrels. In Experiment 2, we test if territoriality motivates tactile interactions by presenting crows with taxidermy crows prepared to look either dead or upright and life-like. In Experiment 1, we find that crows are significantly less likely to make contact but more likely to alarm call and recruit other birds in response to dead crows than to dead pigeons and squirrels. In addition, we find that aggressive and sexual encounters with dead crows are seasonally biased. These findings are inconsistent with feeding or information acquisition-based motivation. In Experiment 2, we find that crows rarely dive-bomb and more often alarm call and recruit other crows to dead than to life-like crows, behaviours inconsistent with responses given to live intruders. Consistent with a danger response hypothesis, our results show that alarm calling and neighbour recruitment occur more frequently in response to dead crows than other stimuli, and that touching dead crows is atypical. Occasional contacts, which take a variety of aggressive and sexual forms, may result from an inability to mediate conflicting stimuli.
This article is part of the theme issue ‘Evolutionary thanatology: impacts of the dead on the living in humans and other animals’.
Evolutionary thanatology benefits from broad taxonomic comparisons of non-human animals' responses to death. Furthermore, exploring the sensory and cognitive bases of these responses promises to allow classification of the underlying mechanisms on a spectrum from phylogenetically ancient to more derived traits. We draw on studies of perception and cognition in invertebrate and vertebrate taxa (with a focus on arthropods, corvids, proboscids, cetaceans and primates) to explore the cues that these animals use to detect life and death in others, and discuss proximate and ultimate drivers behind their capacities to do so. Parallels in thanatological behaviour exhibited by the last four taxa suggest similar sensory–cognitive processing rules for dealing with corpses, the evolution of which may have been driven by complex social environments. Uniting these responses is a phenomenon we term ‘animacy detection malfunction’, whereupon the corpse, having both animate and inanimate attributes, creates states of fear/curiosity manifested as approach/avoidance behaviours in observers. We suggest that integrating diverse lines of evidence (including the ‘uncanny valley’ effect originating from the field of robotics) provides a promising way to advance the field, and conclude by proposing avenues for future research.
This article is part of the theme issue ‘Evolutionary thanatology: impacts of the dead on the living in humans and other animals’.
Information about responses to death in nonhuman primates is important for evolutionary thanatology. This paper reviews the major causes of death in chimpanzees, and how these apes respond to cues related to dying and death. Topics covered include disease, human activities, predation, accidents and intra-species aggression and cannibalism. Chimpanzees also kill and sometimes eat other species. It is argued that, given their cognitive abilities, their experiences of death in conspecifics and other species are likely to equip chimpanzees with an understanding of death as cessation of function and irreversible. Whether they might understand that death is inevitable—including their own death, and biological causes of death is also discussed. As well as gathering more fundamental information about responses to dying and death, researchers should pay attention to possible cultural variations in how great apes deal with death.
This article is part of the theme issue ‘Evolutionary thanatology: impacts of the dead on the living in humans and other animals’.
Societies, including those of humans, have evolved multiple ways of dealing with death across changing circumstances and pressures. Despite many studies focusing on specialized topics, for example necrophoresis in eusocial insects, mortuary activities in early human societies, or grief and mourning in bereavement, there has been little attempt to consider these disparate research endeavours from a broader evolutionary perspective. Evolutionary thanatology does this by adopting an explicit evolutionary stance for studies of death and dying within the sociological, psychological and biological disciplines. The collection of papers in this themed issue demonstrates the value of this approach by describing what is known about how various nonhuman species detect and respond to death in conspecifics, how problems of disposing of the dead have evolved in human societies across evolutionary time and also within much shorter time frames, how human adults' understanding of death develops, and how it is ultimately reflected in death-related language. The psychological significance and impact of death is clearly seen in some species' grief-like reactions to the loss of attachment figures, and perhaps uniquely in humans, the existence of certain psychological processes that may lead to suicide. Several research questions are proposed as starting points for building a more comprehensive picture of the ontogeny and phylogeny of how organisms deal with death.
This article is part of the theme issue ‘Evolutionary thanatology: impacts of the dead on the living in humans and other animals’.
Some aquatic mammals appear to care for their dead, whereas others abandon their live offspring when conditions are unfavourable. This incredible variety in behaviours suggests the importance of comparing and contrasting mechanisms driving death-related behaviours among these species. We reviewed 106 cases of aquatic mammals (81 cetaceans and 25 non-cetaceans) reacting to a death event, and extrapolated ‘participant’ (age class, sex, relationship and decomposition) and ‘social’ characteristics (escorting, calf dependence, alloparental care, herding and dispersal patterns) from published and unpublished literature. A multiple correspondence analysis (MCA) was performed to explore the relationships between these characteristics and death-related behaviours, with species clustered based on MCA scores. Results showed that both cetaceans and non-cetaceans react to death but in different ways. Non-cetaceans, characterized by a short maternal investment, were observed to protect the dead (defending it from external attacks), while cetaceans spent much longer with their offspring and display carrying (hauling, spinning, mouthing with the carcass and diving with it) and breathing-related (lifting and sinking the carcass) activities with the dead generally in association with other conspecifics. Our work emphasizes the need of increased documentation of death-related cases around the world to improve our understanding of aquatic mammals and their responses to death.
This article is part of the theme issue ‘Evolutionary thanatology: impacts of the dead on the living in humans and other animals’.
Maternal cannibalism, whereby a mother consumes her own offspring, occurs in various animal taxa and is commonly explained by nutritional stress or environmental pressures. It is rare in nonhuman primates and is considered an aberrant behavior only observed under high-stress conditions. It was therefore surprising when, in the first reported case of cannibalism in wild bonobos, a mother consumed part of the dead infant at LuiKotale. Here we report two more cases of maternal cannibalism by wild bonobos at two different study sites, Wamba and Kokolopori. The dead infants’ mothers participated in the cannibalism in both cases. At Kokolopori, although the mother did consume part of the carcass, it was held and shared by another dominant female. At Wamba, the mother was a dominant female within the community and was the primary consumer of the carcass. In both cases, cannibalism resembled other meat-eating events, with the dominant female controlling meat consumption. Infanticide was not observed in either case, but its occurrence could not be ruled out. Although rare, the occurrence of maternal cannibalism at three different study sites suggests that this may represent part of the behavioral repertoire of bonobos, rather than an aberrant behavior.
The wounding or death of a conspecific has been shown to elicit varied behavioral responses throughout thanatology. Recently, a number of reports have presented contentious evidence of epimeletic behavior towards the dying and dead among non-human animals, a behavioral trait previously considered uniquely human. Here, we report on the behavioral responses of Barbary macaques, a social, non-human primate, to the deaths of four group members (one high-ranking adult female, one high-ranking adult male, one juvenile male, and one female infant), all caused by road traffic accidents. Responses appeared to vary based on the nature of the death (protracted or instant) and the age class of the deceased. Responses included several behaviors with potential adaptive explanations or consequences. These included exploration, caretaking (guarding, carrying, and grooming), and proximity to wounded individuals or corpses, and immediate as well as longer-lasting distress behaviors from other group members following death, all of which have been reported in other non-human primate species. These observations add to a growing body of comparative evolutionary analysis of primate thanatology and help to highlight the multifaceted impacts of human-induced fatalities on an endangered and socially complex primate.
frequently visited by elephants. At the time of his death, approximately 60 elephants attended, but they dispersed quickly into the surrounding bush during the euthanasia process. Two elephant researchers were present at the time of death, and opportunistically observed the response of the returning elephants to the fresh carcass. From 15 minutes after death until 2.5 hours later, we conducted continuous recording with the carcass as the focus. All contact behaviours and chemosensory investigations were recorded. Contact behaviours included using the torso or trunk to contact various parts of the receiver, including the head and body. The chemosensory behaviours included 'check' (touching the tip of the trunk to a substance), and 'flehmen' (touching the trunk tip to the openings of the vomeronasal organ) (Schulte and Rasmussen 1999), as well as more general trunk touches to the body and wounds. A scan sample also was conducted every minute, noting the proximity and identity of the five nearest individuals to the carcass. Thirty elephants approached the carcass over the 2.5 hours, with 22 animals being scored as one of the five nearest neighbours and 15 animals performing a contact or chemosensory behaviour. Of the 307 total behaviours noted, three animals performed 83.1% of the observed behaviours, while the other 12 individuals were below 5%. The three elephants that performed the majority of behaviours were a male of 21 years (37%), an 18 year old male (34%), and a male of 10 years (13%). These three males also were recorded as the nearest neighbour for 80% of the observation time (30%, 35% and 14%, respectively). These three males were unrelated (maternally) to the dead elephant. Elephants often use the trunk tip chemosensory behaviours of check and flehmen to investigate the environment and to assess the status of conspecifics (Schulte et al. 2005). As with many species, elephants In the wild, African savannah elephants (Loxodonta africana) have been observed to touch dead elephants and manipulate the bones. Cynthia Moss (1988) writes of how elephants in Amboseli regularly investigated elephant carcasses, smelling and touching the bones and skull. In a controlled study, researchers from Am-boseli found that elephants demonstrated higher interest in the skulls of elephants than those of other species (McComb et al. 2006). At Samburu National Reserve in Kenya, Douglas-Hamilton et al. (2006) observed the death of a matriarch and the subsequent responses by her own family and four unrelated families to the carcass. In Addo Elephant National Park, elephants of a variety of ages also express interest in elephant bones, touching them and turning them over (pers. obs.). This examination of elephant bones and ivory indicates that elephants show an elevated level of interest in conspecifics over other dead animals. Elephants do not seem to express special interest in dead kin but rather they appear to have a generalized response to injured, dying and deceased conspecifics (Douglas-Hamilton et al. 2006, McComb et al. 2006). The present study reports the behaviour of a group of elephants in response to a euthanized adult male elephant that suffered severe wounds inflicted by a conspecific male. Most of the observations from previous studies have been on females and female group members. In the present study, of special interest is the behaviour of unrelated, conspecific males to this carcass. Addo Elephant National Park, in the Eastern Cape of South Africa, had a population of 350 elephants during this study. All individuals in the park were known, and the matrilineal family trees have been reconstructed since the park was founded in 1931 (Whitehouse and Hall-Martin 2000). In January 2005, a 41 year old adult male received serious wounds to the head and ears in a conflict with another elephant. He was subsequently euthanized by park staff within 50 m of a waterhole
Spatial memory facilitates resource acquisition where resources are patchy, but how it influences movement behaviour of wide-ranging species remains to be resolved. We examined African elephant spatial memory reflected in movement decisions regarding access to perennial waterholes. State-space models of movement data revealed a rapid, highly directional movement behaviour almost exclusively associated with visiting perennial water. Behavioural change point (BCP) analyses demonstrated that these goal-oriented movements were initiated on average 4.59 km, and up to 49.97 km, from the visited waterhole, with the closest waterhole accessed 90% of the time. Distances of decision points increased when switching to different waterholes, during the dry season, or for female groups relative to males, while selection of the closest waterhole decreased when switching. Overall, our analyses indicated detailed spatial knowledge over large scales, enabling elephants to minimize travel distance through highly directional movement when accessing water. We discuss the likely cognitive and socioecological mechanisms driving these spatially precise movements that are most consistent with our findings. By applying modern analytic techniques to high-resolution movement data, this study illustrates emerging approaches for studying how cognition structures animal movement behaviour in different ecological and social contexts.
© 2015 The Author(s) Published by the Royal Society. All rights reserved.
Long-term social memory is important, because it is an ecologically relevant test of cognitive capacity, it helps us understand which social relationships are remembered and it relates two seemingly disparate disciplines: cognition and sociality. For dolphins, long-term memory for conspecifics could help assess social threats as well as potential social or hunting alliances in a very fluid and complex fission-fusion social system, yet we have no idea how long dolphins can remember each other. Through a playback study conducted within a multi-institution dolphin breeding consortium (where animals are moved between different facilities), recognition of unfamiliar versus familiar signature whistles of former tank mates was assessed. This research shows that dolphins have the potential for lifelong memory for each other regardless of relatedness, sex or duration of association. This is, to my knowledge, the first study to show that social recognition can last for at least 20 years in a non-human species and the first large-scale study to address long-term memory in a cetacean. These results, paired with evidence from elephants and humans, provide suggestive evidence that sociality and cognition could be related, as a good memory is necessary in a fluid social system.
Two different dead bottlenose dolphins were observed in separate years with attending male bottlenose dolphins. In 2000, an adult female bottlenose dolphin body was identified ~150 m from shore in ~8 m depth with two attendant adult male bottlenose dolphins. These males were sighted in proximity with the carcass for two days. In 2001, a sub-adult male bottlenose dolphin was observed ~75 m from shore in ~6 m depth with more than two dozen attendant sub-adult and adult male dolphins and two adult female bottlenose dolphins. Attendant dolphins approached the dead male body and engaged in inquisitive behaviours (e.g., echolocation, head-scanning, nudging, Dudzinski, 1998) at the genital region and chest repeatedly. After 17 min of initial observation, attendant males exhibited erections when positioned within 0.5 m of the carcass. Documented behaviours of attendant dolphins in 2000 correspond with mate-guarding activity previously reported for bottlenose dolphins. Several alternative explanations (e.g., dominance, competition) for behaviours recorded during attendance of the sub-adult male carcass are discussed.
Necrophobic behaviors, defined as the avoidance of dead or injured conspecifics, have been formally documented in insects and aquatic organisms. It is plausible that such avoidance has been selected for by the risks of predation and disease that are associated with the presence of cadavers, and that necrophobic behaviors may be present across a variety of taxa. We demonstrate the avoidance of a house mouse (Mus musculus) cadaver by small mammals visiting experimental food trays, and by male and female house mice in a Y-maze exploration paradigm. In addition, we present individual differences in the responses of house mice to a house mouse cadaver. Further, we propose potential applications for the study of necrophobic behavior in improving wildlife management practices and models in disease ecology.
A 21-month individual identification project on the Samburu and Buffalo Springs National Reserves' elephant population was conducted between November 1997 and July 1999. The free ranging population, of at least 767 elephants, which relied heavily on areas outside the reserves, was individually identified. The numbers of elephants observed per day fluctuated but were greater during dry periods then wet. However, the sizes of aggregations were greater during wet periods. Preliminary investigation suggested that the population could be divided into two groups, which were designated resident and non-resident family units. The groups comprised approximately equal numbers of cows and calves, but temporally had different reserve use patterns and calving peaks. The daily numbers of males and musth males were correlated with numbers of females. The reserves appeared to be a focal area for calving, indicating that the study area was of reproductive importance to the population. Demographic data indicated a female biased population sex ratio, with over twice the number of mature females to males. The observed sex skew was greatest for older age classes, and the density of musth bulls in the study area was low. The population was affected by poaching. Continued monitoring will assist conservation efforts by alerting authorities of major demographic or range use changes.
The structure of dominance relationships among individuals in a population is known to influence their fitness, access to
resources, risk of predation, and even energy budgets. Recent advances in global positioning system radio telemetry provide
data to evaluate the influence of social relationships on population spatial structure and ranging tactics. Using current
models of socio-ecology as a framework, we explore the spatial behaviors relating to the maintenance of transitive (i.e.,
linear) dominance hierarchies between elephant social groups despite the infrequent occurrence of contests over resources
and lack of territorial behavior. Data collected from seven families of different rank demonstrate that dominant groups disproportionately
use preferred habitats, limit their exposure to predation/conflict with humans by avoiding unprotected areas, and expend less
energy than subordinate groups during the dry season. Hence, our data provide strong evidence of rank derived spatial partitioning
in this migratory species. These behaviors, however, were not found during the wet season, indicating that spatial segregation
of elephants is related to resource availability. Our results indicate the importance of protecting preexisting social mechanisms
for mitigating the ecological impacts of high density in this species. This analysis provides an exemplar of how behavioral
research in a socio-ecological framework can serve to identify factors salient to the persistence and management of at risk
species or populations.
In this paper, we investigate the formation and function of the multilevelled, fission–fusion social structure in a free-ranging African elephant, Loxodonta africana, population. We quantitatively identified the existence of four social tiers by using cluster analysis on individual association data. We assessed the effects of season and study period on social structuring and levels of cohesion within and among social units. We found that second-tier units, potentially the equivalent of the ‘family’, were stable across seasonal periods but the number of units increased as the study progressed and the population grew. It appears that these units were sufficiently small not to be influenced by ecologically related factors, such as resource competition, that might otherwise lead to them splitting. On the other hand, third- and fourth-tier units were significantly affected by season in a way that suggests a trade-off between ecological costs (e.g. from resource competition) and different social and ecological benefits (e.g. from predator defence, territoriality, knowledge sharing and rearing of young). Age structure also appeared to influence this multitiered social organization. The size of second-tier social units was significantly affected by the age of matriarchs: units lead by matriarchs likely to be grandmothers (i.e. females 35 years and older) were significantly larger than those lead by younger matriarchs. We present a conceptual framework for understanding the emergence of multiple-tier social structure from interactions driven by socioecological processes. This study is the first to use rigorous quantitative methods to statistically show the existence of four hierarchical tiers of social organization in a nonhuman animal. Additionally, our results elucidate the role that ecological processes play in producing complex social structures.
Dolphins are well known for their exquisite echolocation abilities, which enable them to detect and discriminate prey species and even locate buried prey. While these skills are widely used during foraging, some dolphins use tools to locate and extract prey. In the only known case of tool use in free-ranging cetaceans, a subset of bottlenose dolphins (Tursiops sp.) in Shark Bay, Western Australia habitually employs marine basket sponge tools to locate and ferret prey from the seafloor. While it is clear that sponges protect dolphins' rostra while searching for prey, it is still not known why dolphins probe the substrate at all instead of merely echolocating for buried prey as documented at other sites. By 'sponge foraging' ourselves, we show that these dolphins target prey that both lack swimbladders and burrow in a rubble-littered substrate. Delphinid echolocation and vision are critical for hunting but less effective on such prey. Consequently, if dolphins are to access this burrowing, swimbladderless prey, they must probe the seafloor and in turn benefit from using protective sponges. We suggest that these tools have allowed sponge foraging dolphins to exploit an empty niche inaccessible to their non-tool-using counterparts. Our study identifies the underlying ecological basis of dolphin tool use and strengthens our understanding of the conditions that favor tool use and innovation in the wild.
Patterns in the association of individuals can shed light on the underlying conditions and processes that shape societies. Here we characterize patterns of association in a population of wild Asian Elephants at Uda Walawe National Park in Sri Lanka. We observed 286 individually-identified adult female elephants over 20 months and examined their social dynamics at three levels of organization: pairs of individuals (dyads), small sets of direct companions (ego-networks), and the population level (complete networks).
Corroborating previous studies of this and other Asian elephant populations, we find that the sizes of elephant groups observed in the field on any particular day are typically small and that rates of association are low. In contrast to earlier studies, our longitudinal observations reveal that individuals form larger social units that can be remarkably stable across years while associations among such units change across seasons. Association rates tend to peak in dry seasons as opposed to wet seasons, with some cyclicity at the level of dyads. In addition, we find that individuals vary substantially in their fidelity to companions. At the ego-network level, we find that despite these fluctuations, individuals associate with a pool of long-term companions. At the population level, social networks do not exhibit any clear seasonal structure or hierarchical stratification.
This detailed longitudinal study reveals different social dynamics at different levels of organization. Taken together, these results demonstrate that low association rates, seemingly small group sizes, and fission-fusion grouping behavior mask hidden stability in the extensive and fluid social affiliations in this population of Asian elephants.
Climate change in Africa is expected to lead to a higher occurrence of severe droughts in semi-arid and arid ecosystems. Understanding how animal populations react to such events is thus crucial for addressing future challenges for wildlife management and conservation. We explored how gender, age, mother's experience and family group characteristics determined calf survival in an elephant population during a severe drought in Tanzania in 1993. Young males were particularly sensitive to the drought and calf loss was higher among young mothers than among more experienced mothers. We also report high variability in calf mortality between different family groups, with family groups that remained in the National Park suffering heavy calf loss, compared with the ones that left the Park. This study highlights how severe droughts can dramatically affect early survival of large herbivores and suggests that extreme climatic events might act as a selection force on vertebrate populations, allowing only individuals with the appropriate behaviour and/or knowledge to survive.
An important area of biology involves investigating the origins in animals of traits that are thought of as uniquely human. One way that humans appear unique is in the importance they attach to the dead bodies of other humans, particularly those of their close kin, and the rituals that they have developed for burying them. In contrast, most animals appear to show only limited interest in the carcasses or associated remains of dead individuals of their own species. African elephants (Loxodonta africana) are unusual in that they not only give dramatic reactions to the dead bodies of other elephants, but are also reported to systematically investigate elephant bones and tusks that they encounter, and it has sometimes been suggested that they visit the bones of relatives. Here, we use systematic presentations of object arrays to demonstrate that African elephants show higher levels of interest in elephant skulls and ivory than in natural objects or the skulls of other large terrestrial mammals. However, they do not appear to specifically select the skulls of their own relatives for investigation so that visits to dead relatives probably result from a more general attraction to elephant remains.
Among terrestrial mammals, elephants share the unique status, along with humans and great apes, of having large brains, being long-lived and having offspring that require long periods of dependency. Elephants have the largest brains of all terrestrial mammals, including the greatest volume of cerebral cortex. In contrast to what one might expect from such a large-brained species, the performance of elephants in cognitive feats, such as tool use, visual discrimination learning and tests of "insight" behavior, is unimpressive in comparison to the performance by chimpanzees and, of course, humans. Where elephants do seem to excel is in long-term, extensive spatial-temporal and social memory. In addition, elephants appear to be somewhat unique among non-human species in their reactions to disabled and deceased conspecifics, exhibiting behaviors that are mindful of "theory-of-mind" phenomena. Information gleaned from studies on the neural cytoarchitecture of large brains reveals that the neurons of the cerebral cortex of elephants are much less densely populated than in large-brained primates. The interactions between cortical neurons would appear to be more global and less compartmentalized into local areas, and cortical information processing slower, than in great apes and humans. Although focused neural cytoarchitecture studies on the elephant are needed, this comparative perspective on the cortical neural cytoarchitecture appears to relate to differences in behavior between elephants and their primate counterparts.
Monitoring the location of conspecifics may be important to social mammals. Here, we use an expectancy-violation paradigm to test the ability of African elephants (Loxodonta africana) to keep track of their social companions from olfactory cues. We presented elephants with samples of earth mixed with urine from female conspecifics that were either kin or unrelated to them, and either unexpected or highly predictable at that location. From behavioural measurements of the elephants' reactions, we show that African elephants can recognize up to 17 females and possibly up to 30 family members from cues present in the urine-earth mix, and that they keep track of the location of these individuals in relation to themselves.
Social environments are fundamental to fitness in many species. In disrupted societies, the loss of important partners may alter social environments for surviving individuals. African elephants, Loxodonta africana, have experienced age-selective mortality linked to the ivory trade, and the resulting social costs for surviving young elephants are unknown. In this study, we followed orphaned female elephants and nonorphaned counterparts in Kenya's Samburu and Buffalo Springs National Reserves to elucidate whether orphaning and related dispersal behaviour incur social costs. There were clear social differences between orphans and nonorphans, most notably in that orphans tended to receive more aggression than nonorphans. Dispersal from natal groups was a behaviour found exclusively among orphans. Differences in social environments of orphans that remained in their natal groups and those that dispersed were also found in the form of dispersed orphans receiving more aggression while feeding than those that remained in their natal group. Our results suggest that orphaning in elephants is associated with social costs, and that these costs are amplified for orphans that disperse from their natal groups. Future research should identify the relationship between the social costs of being an orphan and fitness, which may be important to the recovery of populations affected by the ivory trade and other forms of disruption.
Network resilience to perturbation is fundamental to functionality in systems ranging from synthetic communication networks to evolved social organization [1]. While theoretical work offers insight into causes of network robustness, examination of natural networks can identify evolved mechanisms of resilience and how they are related to the selective pressures driving structure. Female African elephants (Loxodonta africana) exhibit complex social networks with node heterogeneity in which older individuals serve as connectivity hubs [2, 3]. Recent ivory poaching targeting older elephants in a well-studied population has mirrored the targeted removal of highly connected nodes in the theoretical literature that leads to structural collapse [4, 5]. Here we tested the response of this natural network to selective knockouts. We find that the hierarchical network topology characteristic of elephant societies was highly conserved across the 16-year study despite ∼70% turnover in individual composition of the population. At a population level, the oldest available individuals persisted to fill socially central positions in the network. For analyses using known mother-daughter pairs, social positions of daughters during the disrupted period were predicted by those of their mothers in years prior, were unrelated to individual histories of family mortality, and were actively built. As such, daughters replicated the social network roles of their mothers, driving the observed network resilience. Our study provides a rare bridge between network theory and an evolved system, demonstrating social redundancy to be the mechanism by which resilience to perturbation occurred in this socially advanced species.
C. J. Moss, H. Croze, P. C. Lee (eds.). 2011. The Amboseli Elephants: A Long-term Perspective on a Long-lived Mammal. University of Chicago Press, Chicago, Illinois, 383 pp. ISBN-13 978-0-226-54223-2 and ISBN-10 0-226-54223-8, price (hardbound), $65.00.
A 38-year program of research on a single population, the Amboseli Elephant Research Project, stands as a pinnacle example of mammalogy for its length and depth of investigation. This study, widely known through the scientific and popular writing of Cynthia Moss and Joyce Poole, is now put into full scientific context in this edited multiauthor volume that was 12 years in the making, or as the authors point out, “the gestation period of 5–6 elephant calves.”
What the authors have achieved is not just a reference on the Amboseli population, but a compendium of biological, ecological, and natural history knowledge on elephants. By comparison, all the other great books on elephant behavior read more like travelogues or biographies. Be warned, readers looking for the common coffee table book with large glossy color photos and field anecdotes will be sorely disappointed. Appealingly, this book instead focuses on providing a well-organized overview of the Amboseli system and the elephants within it by summarizing previously published research findings (with updated information), as well as novel information not published in the peer-reviewed literature during this long-term project.
The book is organized into 5 parts. In Part 1, the editors set the background for their subsequent detailed sections on elephant behavior. Harvey Croze and Keith Lindsay provide a background chapter devoted to the biotic and abiotic conditions of Amboseli and how those conditions changed over time. Kadzo Kangwana and Christine Brown-Nunez provide an intriguing and detailed account of how Maasai communities surrounding the park are shifting toward a more sedentary agricultural lifestyle that is incorporated into the commercial economy. Treated objectively in Part 1, the increasing importance of human land use and interactions with elephants is a recurrent theme throughout the book, and is revisited from the perspective of Amboseli Elephant Research Project proponents in …
The study of animal death is poised to blossom into an exciting new interdisciplinary field-and one with profound relevance for bioethics. Areas of interest include the biology and evolution of death-related behavior in nonhuman animals, as well as human social, psychological, cultural, and moral attitudes toward and practices related to animal death. In this paper, I offer a brief overview of what we know about death-related behavior in animals. I will then sketch some of the bioethical implications of this emerging field of research.
The extent to which elephants hold behavioural traits in common with human beings is relevant to the ethics of how we treat them. Observations show that elephants, like humans, are concerned with distressed or deceased individuals, and render assistance to the ailing and show a special interest in dead bodies of their own kind. This paper reports helping and investigative behaviour of different elephants and their families towards a dying and deceased matriarch. We make use of long-term association records, GPS tracking data and direct observations. Records made around the time of death, shows that the helping behaviour and special interest exhibited was not restricted to closely related kin. The case is made that elephants, like human beings, can show compassionate behaviour to others in distress. They have a general awareness and curiosity about death, as these behaviours are directed both towards kin and non-related individuals.
Mounting evidence from species as diverse as cats and dolphins indicates that humans are not the only species that grieves over the loss of loved ones
We report morphological data on brains of four African, Loxodonta africana, and three Asian elephants, Elephas maximus, and compare findings to literature. Brains exhibit a gyral pattern more complex and with more numerous gyri than in primates, humans included, and in carnivores, but less complex than in cetaceans. Cerebral frontal, parietal, temporal, limbic, and insular lobes are well developed, whereas the occipital lobe is relatively small. The insula is not as opercularized as in man. The temporal lobe is disproportionately large and expands laterally. Humans and elephants have three parallel temporal gyri: superior, middle, and inferior. Hippocampal sizes in elephants and humans are comparable, but proportionally smaller in elephant. A possible carotid rete was observed at the base of the brain. Brain size appears to be related to body size, ecology, sociality, and longevity. Elephant adult brain averages 4783g, the largest among living and extinct terrestrial mammals; elephant neonate brain averages 50% of its adult brain weight (25% in humans). Cerebellar weight averages 18.6% of brain (1.8 times larger than in humans). During evolution, encephalization quotient has increased by 10-fold (0.2 for extinct Moeritherium, ∼2.0 for extant elephants). We present 20 figures of the elephant brain, 16 of which contain new material. Similarities between human and elephant brains could be due to convergent evolution; both display mosaic characters and are highly derived mammals. Humans and elephants use and make tools and show a range of complex learning skills and behaviors. In elephants, the large amount of cerebral cortex, especially in the temporal lobe, and the well-developed olfactory system, structures associated with complex learning and behavioral functions in humans, may provide the substrate for such complex skills and behavior.
Explored the tool-use potentials of captive African and Asian elephants,
Loxodonta africana and
Elephus maximus, respectively, and wild African elephants. The captive Ss showed 11.6 (Asian) and 22.8 (African) acts of tool use per hour, whereas wild Ss used tools at a frequency of 1.45 acts/hour. With respect to diversity, the captive Ss performed 21 types of tool use, and these tool-using behavior patterns occurred in 3 major functional contexts: body care, aggression, and an ambiguous context. Wild Ss performed 9 types of tool use, which occurred in the same 3 contexts as well as in the context of feeding another. The variability observed in tool-use behavior suggests that, in elephants, tool use may serve as adaptations enabling these furless, large-bodied tropical mammals to cope with ectoparasites and thermoregulation. (PsycINFO Database Record (c) 2012 APA, all rights reserved)
Research on acoustic communication has often focused on signalling between territorial individuals or static neighbouring groups. Under these circumstances, receivers have the opportunity to learn to recognize the signals only of the limited number of conspecifics with which they are in auditory contact. In some mammals, however, social units move freely with respect to one another and range widely, providing individuals with opportunities to learn to recognize the signals of a wide range of conspecifics in addition to those of their immediate neighbours. We conducted playback experiments on African elephants,Loxodonta africana , in Amboseli National Park, Kenya, to determine the extent to which adult female elephants, which have a highly fluid social system, can recognize others in the population through infrasonic contact calls. Female elephants could distinguish the calls of female family and bond group members from those of females outside of these categories; moreover, they could also discriminate between the calls of family units further removed than bond group members, on the basis of how frequently they encountered them. We estimated that subjects would have to be familiar with the contact calls of a mean of 14 families in the population (containing around 100 adult females in total), in order to perform these discriminations. Female elephants thus appear to have unusually extensive networks of vocal recognition, which may prove to be typical of long-lived species that have both fluid social systems and the means for long-distance vocal communication.
The forests surrounding Bossou, Guinea, are home to a small, semi-isolated chimpanzee community studied for over three decades [1]. In 1992, Matsuzawa [2] reported the death of a 2.5-year-old chimpanzee (Jokro) at Bossou from a respiratory illness. The infant's mother (Jire) carried the corpse, mummified in the weeks following death, for at least 27 days. She exhibited extensive care of the body, grooming it regularly, sharing her day- and night-nests with it, and showing distress whenever they became separated. The carrying of infants' corpses has been reported from a number of primate species, both in captivity and the wild [3-7] - albeit usually lasting a few days only - suggesting a phylogenetic continuity for a behavior that is poignant testament to the close mother-infant bond which extends across different primate taxa. In this report we recount two further infant deaths at Bossou, observed over a decade after the original episode but with striking similarities.
Primates undoubtedly have impressive abilities in perceiving, recognizing, manipulating, and predicting other individuals, but only great apes seem to recognize the cognitive basis of manipulative and cooperative tactics or the concept of self. None of these abilities is unique to primates. We distinguish (1) a package of quantitative advantages in social sophistication, perhaps based on more efficient memory, in which neocortical enlargement is associated with the challenge of social living; from (2) a qualitative difference in understanding, whose taxonomic distribution--including several distantly related species, including birds--does not point to an evolutionary origin in social challenges and may instead relate to a need to acquire novel ways of dealing with the physical world. The ability of great apes to learn new manual routines by parsing action components may have driven their qualitatively greater social skill, suggesting that strict partition of physical and social cognition is likely to be misleading.
Cynthia Moss has studied the elephants in Kenya's Amboseli National Park for over twenty-seven years. Her long-term research has revealed much of what we now know about these complex and intelligent animals. Here she chronicles the lives of the members of the T families led by matriarchs Teresia, Slit Ear, Torn Ear, Tania, and Tuskless. With a new afterword catching up on the families and covering current conservation issues, Moss's story will continue to fascinate animal lovers. "One is soon swept away by this 'Babar' for adults. By the end, one even begins to feel an aversion for people. One wants to curse human civilization and cry out, 'Now God stand up for the elephants!'"âChristopher Lehmann-Haupt, New York Times "Moss speaks to the general reader, with charm as well as scientific authority. . . . [An] elegantly written and ingeniously structured account." âRaymond Sokolov, Wall Street Journal "Moss tells the story in a style so conversational . . . that I felt like a privileged visitor riding beside her in her rickety Land-Rover as she showed me around the park." âSarah Blaffer Hrdy, New York Times Book Review "A prose-poem celebrating a species from which we could learn some moral as well as zoological lessons." âChicago Tribune
In order to identify the conditions that favored the flourishing of primate tool use into hominid technology, we examine inter- and intraspecific variation in manufacture and use of tools in extant nonhuman primates, and develop a model to account for their distribution. We focus on tools used in acquiring food, usually by extraction. Any model for the evolution of the use of feeding tools must explain why tool use is found in only a small subset of primate species, why many of these species use tools much more readily in captivity, why routine reliance on feeding tools is found in only two species of ape, and why there is strong geographic variation within these two species. Because ecological factors alone cannot explain the distribution of tool use in the wild, we develop a model that focuses on social and cognitive factors affecting the invention and transmission of tool-using skills. The model posits that tool use in the wild depends on suitable ecological niches (especially extractive foraging) and the manipulative skills that go with them, a measure of intelligence that enables rapid acquisition of complex skills (through both invention and, more importantly, observational learning), and social tolerance in a gregarious setting (which facilitates both invention and transmission). The manipulative skills component explains the distribution across species of the use of feeding tools, intelligence explains why in the wild only apes are known to make and use feeding tools routinely, and social tolerance explains variation across populations of chimpanzees and orangutans. We conclude that strong mutual tolerance was a key factor in the explosive increase in technology among hominids, probably intricately tied to a lifestyle involving food sharing and tool-based processing or the acquisition of large, shareable food packages.
We report morphological data on brains of four African, Loxodonta africana, and three Asian elephants, Elephas maximus, and compare findings to literature. Brains exhibit a gyral pattern more complex and with more numerous gyri than in primates, humans included, and in carnivores, but less complex than in cetaceans. Cerebral frontal, parietal, temporal, limbic, and insular lobes are well developed, whereas the occipital lobe is relatively small. The insula is not as opercularized as in man. The temporal lobe is disproportionately large and expands laterally. Humans and elephants have three parallel temporal gyri: superior, middle, and inferior. Hippocampal sizes in elephants and humans are comparable, but proportionally smaller in elephant. A possible carotid rete was observed at the base of the brain. Brain size appears to be related to body size, ecology, sociality, and longevity. Elephant adult brain averages 4783 g, the largest among living and extinct terrestrial mammals; elephant neonate brain averages 50% of its adult brain weight (25% in humans). Cerebellar weight averages 18.6% of brain (1.8 times larger than in humans). During evolution, encephalization quotient has increased by 10-fold (0.2 for extinct Moeritherium, approximately 2.0 for extant elephants). We present 20 figures of the elephant brain, 16 of which contain new material. Similarities between human and elephant brains could be due to convergent evolution; both display mosaic characters and are highly derived mammals. Humans and elephants use and make tools and show a range of complex learning skills and behaviors. In elephants, the large amount of cerebral cortex, especially in the temporal lobe, and the well-developed olfactory system, structures associated with complex learning and behavioral functions in humans, may provide the substrate for such complex skills and behavior.
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On the ecology and behaviour of the African elephant
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Reproductive tactics of male African savannah elephants (Loxodonta africana)
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The natural history of the African Elephant
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The Amboseli elephants: a long-term perspective on a long-lived mammal
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Large brains and cognition: where do elephants fit in?
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