Reactions to dying and dead conspecifics have been observed in many non-human animals. Elephants, particularly African elephants, are thought to have an awareness of the death of their conspecifics, as they show compassionate behaviour towards others in distress. However, there is a paucity of scientific documentation on thanatological responses displayed by Asian elephants. Here, we report three detailed, directly observed cases of free-ranging Asian elephants (Elephas maximus) responding to dying and dead conspecifics. Behavioural responses were recorded opportunistically and described as pre-, peri- and post-mortem phases based on the status of the individual before, near or after its death. In all three observations, elephants showed approach and exploratory (sniffing and inspecting) behaviours, and epimeletic or helping (physically supporting dying calves) in pre- and peri-mortem phases. We also recorded high-frequency vocalizations (trumpets) by an adult female in the presence of a dying calf. Our observations indicate that, like their African counterparts, Asian elephants might experience distress in response to the death of conspecifics, and may have some awareness of death. This information furthers our understanding of the emotional and cognitive complexities of highly social elephants, and contributes to the growing field of elephant thanatology.
Comprehension of at least some subcomponents of the mature concept of death (inevitability, irreversibility, non-functionality and causality) is no longer held to be uniquely human (Anderson 2016; Gonçalves and Biro 2018). Recently, increasing attention has been paid to understanding both proximate and ultimate causation behind the responses of ‘non-human animals’ to death (Anderson et al. 2018; Gonçalves and Biro 2018). Anecdotal evidence, case reports and scientific documentation on insects, corvids, ungulates, carnivores, cetaceans, sirenians and non-human primates clearly show that many species ‘respond’ to dying and dead individuals (Gonçalves and Biro 2018). As these responses vary across species (representing a wide range of phylogenetically ancient to more advanced traits) and reflect differences in underlying cognitive ability, social organisation, and perceptual biases (Gonçalves and Biro 2018), a comparative thanatological perspective is important in order to better understand the evolutionary bases and development of ‘death awareness’ across taxa.
Death-context behaviours depend on different combinations of olfactory, visual, vocal and tactile cues (also lack of cues; e.g., movement, sounds; Gonçalves and Biro 2018). Confirmed thanatological behaviours include mobbing, transport, removal (particularly in eusocial insects), burial (again, in eusocial insects and also in African elephants), ‘ceremonial gathering’ and cacophonous aggregations (in corvids), alarm calls and other vocalizations (corvids, macaques, African elephants), dead infant carrying (non-human primates and cetaceans), aggression, vigils/guarding, cautious inspection, curiosity/approach, and visitations (African elephant: Douglas-Hamilton et al. 2006; Hawley et al. 2018; Bercovitch 2019; cetaceans: Quintana-Rizzo and Wells 2016; Bearzi et al. 2017; corvids: Iglesias et al. 2012; Swift and Marzluff 2018; dingo: Appleby et al. 2013; eusocial insects: López-Riquelme and Fanjul-Moles 2013; Sun et al. 2013; primates: Anderson 2017; Watson and Matsuzawa 2018; Gonçalves and Carvalho 2019; ungulates: Bercovitch 2012). It has been suggested that species with similar sensory and cognitive abilities (e.g., large brains, highly developed cognitive processing, information-gathering and causal reasoning abilities) show similar responses to the dead (Gonçalves and Biro 2018).
One of these taxa, the extant proboscideans, have complex cognitive abilities, empathy and complex social structures (Sukumar 2003; Bates et al. 2008; Hart et al. 2008). In both African savannah and forest elephants, there are reports of surviving individuals responding to sick, dying or dead elephants by staying near and guarding them (Douglas-Hamilton et al. 2006; Hawley et al. 2018), showing chemosensory (‘check’ and ‘flehmen’) behaviours along with general trunk touches (Merte et al. 2008), gentle kicking, touching and holding the dying or dead by the forefeet and using tusks to lift the carcass (Douglas-Hamilton et al. 2006; McComb et al. 2006; Poole and Granli 2011), revisiting, carrying, covering (with dirt, leaves, branches, etc.), inspecting, and vocalizing (Moss 1988; Poole and Granli 2011; Gonçalves and Biro 2018). McComb et al. (2006) described field experiments in which African savannah elephants (Loxodonta africana) showed greater interest in bones of conspecifics than other species, and in conspecific tusks compared to skulls. Douglas-Hamilton et al. (2006) reported that African elephants showed helping behaviours towards dying conspecifics irrespective of their genetic relatedness. However, there is a notable lack of similar scientific documentation for their proboscidean relatives, the Asian elephants (Elephas maximus).
Having diverged between 9 and 4.2 million years ago (Mya) from the African elephant (genus Loxodonta; Palkopoulou et al. 2018), Asian elephants share several morphological, social and behavioural similarities with their closest extant relatives, but also show some differences. Like their African savannah counterparts, Asian elephants live in fluid, multi-tiered fusion-fission matriarchal societies, with close bonds among members of family units (Sukumar 2003; Nandini et al. 2017); however, they do not show clear hierarchical structures in their societies (de Silva et al. 2017). With respect to their morphology, Asian elephants have relatively smaller height, body mass and ear size. One striking difference between the two species is that female Asian elephants are universally devoid of full-grown tusks, whereas most African female elephants possess tusks; furthermore, many male Asian elephants are also tuskless, a rare condition among male African elephants (Sukumar 2003). Despite these differences, the species share complex cognitive abilities (Bates et al. 2008; Plotnik et al. 2010), so the question arises whether Asian elephants behave similarly to African elephants in death-related contexts.
It is rare to observe the precise moment of death of an elephant in the wild. Concerning Asian elephants, there are unconfirmed anecdotes about “burying” dead or injured humans and other animals (Sukumar 1994), visiting the spot where an elephant died after accidental electrocution in a village, and repeatedly visiting the carcass of familiar conspecifics. One such record was made in the Kaziranga National Park, India, during October 2012 when a female elephant (c. 40 years) was seen carrying her dead calf for several days (Chelliah and Sukumar, unpublished video). Although there are some news reports and videos of Asian elephants responding to injured calves and carrying carcasses, there have been no formal scientific descriptions of thanatological behaviour in Asian elephants.
Here, we report three opportunistic observations of behavioural reactions of free-ranging Asian elephants to dying and dead conspecifics. We categorized the observations to document how Asian elephants react to (1) a dying newborn calf (pre-mortem phase), (2) an injured and dying calf (peri-mortem phase), and (3) a dead adult female (post-mortem phase).
The observations were conducted in Bandipur National Park (BNP, 11.5923°–11.9488°N, 76.2085°–76.8690°E) and the adjoining areas of Mudumalai National Park (MNP, 11.7055°–11.4683°N, 76.3577°–77.0177°E) of the Nilgiri-Mysore-Wyanad Elephant Reserve in southern India. Both reserves have a general elevation range from c. 800–1100 m. The study area shelters one of the high densities of Asian elephants, with BNP having c. 1025 elephants in an area of 906 km2 and MNP having c. 300 elephants in an area of 321 km2 (Synchronized Elephant Population Estimation 2017). With an average annual rainfall of 800–1300 mm in BNP and 700–1800 mm (along an east–west gradient) in MNP, the predominant vegetation types in both national parks are seasonal dry tropical forests, with MNP having a dry thorn forest in the east, dry and moist deciduous forest in the central region, and patches of semi-evergreen forest to the west (Sukumar et al. 2004). The Moyar River in MNP forms one of the perennial sources of water for elephants and other animals. The Kabini reservoir (along the northern boundary of BNP) witnesses a large congregation of elephants during the resource-deficient dry season (March–May).
Opportunistic observations were made by two researchers working independently on long-term monitoring of population dynamics of Asian elephants and stress physiology projects, during fieldwork at two different sites. Elephants’ age classes were estimated through the relative height method (Sukumar 1989). Individual identification, particularly in the case of Sita (an adult female who was first sighted in 1990), was based on morphological features such as ears (shape, presence of holes or cuts, folds, pigmentation and venation patterns), marks or warts on the body, and tail length with tail hair shape (Vidya et al. 2014). Behaviours were categorized as pre-mortem (towards a dying newborn calf), peri-mortem (towards an injured and dying calf) and post-mortem (towards a dead adult female) observations, and were noted as continuous sampling and recorded on camera (two events) including video graphs (one event; Sony HDR-PJ540 Handycam). All three observations were made on foot from a safe and non-intrusive distance of 60–100 m between elephants and observers. Vocalizations were recorded using an Earthworks QTC-50 microphone (omnidirectional, with a frequency response of 3 Hz to 50 kHz) embedded in a Rycote modular windshield connected to a Fostex DC-R 302 recorder and at a sampling rate of 48 kHz. The acoustic properties of the calls (duration and maximum frequency) were calculated using the software Raven Pro 1.5. Below, we present only descriptive accounts; we did not quantify behaviours or perform any statistical analyses.
Three opportunistic observation were recorded of elephants responding to injured, dying or dead conspecifics. Two of the three cases involved adult female-calf pairs and involved epimeletic or helping behaviour towards the injured and dying calf. In all three cases, exploratory and approach/visit behaviours towards the carcass were observed. Detailed behavioural descriptions of each case are provided below, while a detailed summary and the timeline are presented in Table 1 and as Supplementary Material Table 1.
Pre-mortem observations (reactions to a dying newborn calf)
Reactions of an adult female elephant (named Sita) and her daughters (S1: 10–15 years old and S2: 5–10 years old) to her dying newborn calf were observed. The incident occurred on 29 October 2012 at 11.57306°N and 76.61610°E of the MNP. Sita was one of the oldest resident females of MNP (she was first photographed and identified with a calf in 1990 by RS at the MNP). Based on our long-term field observations since 2009, Sita was usually with her two daughters (herd size = 3) and in occasional association with a sub-adult male. Sita was sighted with her two daughters three times in 2012 before the incident took place.
Sita, with her newborn calf, and daughters S1 and S2, were spotted near a highway which traverses the MNP. The three females surrounded the calf with their legs. S1 sniffed the calf, while Sita gently ‘kicked’ with her forelegs in an attempt to make it stand. S1 and S2 joined Sita to help. As Sita kicked gently, S1 stood against the calf, offering her legs as a support. However, the calf failed to stand and fell again to the ground. The gentle kicking and supporting behaviours were repeated by the three females. S1 also touched Sita’s mouth with her trunk, before moving off to graze while Sita and S2 continued supporting the calf (Fig. 1a). Interestingly, Sita and S2 appeared to coordinate their helping actions: Sita used her trunk to grip the calf’s rear end while S2 supported its head with her trunk. However, the calf appeared unable to fully stand up.
Ten min after leaving the scene, S1 returned. Further gentle kicking and supporting with the legs occurred (Fig. 1b). Despite these and its own efforts, the calf repeatedly collapsed while Sita stood over it, keeping the calf between her legs (Fig. 1c, d). The kick-and-support epimeletic behaviour continued for over 3 h. Flehmen-like behaviour was also observed; S1 and S2 sniffed or contacted Sita’s genitals (Fig. 1e). Later, S1 and S2 rested their forehead on each other (Fig. 1f), while standing next to Sita and the calf. All three females touched the calf repeatedly. S1 left again, while Sita and S2 remained by the calf. Spontaneous oozing of milk from Sita’s mammary glands was noted. Apart from encouraging the calf to stand on its feet, Sita and her daughters showed no signs of allomothering or attempts to make it suckle. Up until this point, the total distance moved by Sita was less than 10 m.
Later in the late afternoon, Sita was seen again with the calf, but otherwise alone. The forestry department’s veterinary team provided drinking water for the elephants in a large steel container placed against a nearby tree, but only Sita drank. She showed no signs of aggression as she retreated, and allowed the team to approach the calf. Upon closer examination, the veterinary officials declared the calf as a male and that it had no external injuries. Sita returned to the calf after about 4 min and resumed her attempts to make him stand by gently kicking with her forelegs. The calf eventually reached the dense undergrowth, making it difficult to continue observations. From the spot where she was first sighted about 8 h earlier, Sita had moved only 30 m, and except for the brief period in which she left the calf, the distance between them was less than 1 m. Even though these events occurred next to the highway, no signs of agitation or aggression were observed from Sita and her daughters, and no vocalizations were heard.
The following day, the calf was found dead. The veterinarians could not clearly determine the cause of death, and hypothesized that the calf had died due to dehydration, malnourishment or both. Sita was seen beside her dead calf, but there were no signs of her daughters, S1 and S2. We could not clearly observe Sita’s responses to the carcass due to dense undergrowth; however, over the following week, Sita was often spotted moving and grazing alone, near the spot where her calf died. The entire period of direct observation was 502 min.
Peri-mortem observations (reactions to an injured and dying calf)
The behavioural responses of an adult female (F1: 15–20 years old) to an injured and dying calf (less than 1 year old) were recorded. This incident occurred on 13 March 2017 at 11.59716°N and 76.57493°E of the MNP.
A lone calf (less than 1 year old) was sighted standing near a salt lick inside the MNP. It appeared to be in the poor physical condition and rested its trunk on a dead log. After about 10 min, an adult female (F1) was spotted, standing vigilant almost 40 m away from the calf. A few min later, F1 rushed towards the calf and ‘pushed’ it from behind in an attempt to make it move. The calf could not move much, and turned 180° before collapsing onto its knees. Instantly after the fall, F1 produced a loud ‘trumpet’ call (duration = 1.37 s; maximum frequency = 895.10 Hz; ESM_V1a; Fig. 2a) directed to the calf, followed by another trumpet 17 s later (duration = 0.85 s; maximum frequency = 955.81 Hz; Fig. 2b). She continued to stand beside the calf, and after 1 min since her previous vocalizations, she produced a third trumpet, shorter and of lower frequency (duration = 0.38 s; maximum frequency = 830.49 Hz; Fig. 2c).
The calf stayed in the same position after its fall, moving minimally (slow ear flaps), while F1 touched and sniffed the calf’s genital region. F1 blew saliva on herself (through her trunk) and stood next to the calf, touching it repeatedly. She used her forelegs in alternation to encourage the calf to stand on its feet (ESM_V1b), but without success. She then left the calf and moved 100 m away to graze. During F1’s absence, close inspection revealed that the calf had an open wound on its forehead, probably the result of a tiger (Panthera tigris) attack. After 1 h, F1 ran towards the calf trumpeting (duration and frequency were not measured due to high signal-to-noise ratio). She again stood beside the calf, gently touching, rubbing and pushing it (ESM_V1c). She also kicked the ground to bring the legs of the calf together. However, there was no clear response from the calf.
When the veterinary team arrived, F1 remained vigilant, mock-charged the team and displayed ‘body-guarding’ behaviour with respect to the calf. She ran frantically towards the team, chirping (2–3 times with a bout of 3–4 times each) and then finally made a more determined charge. The people managed to drive F1 away from the calf; however, she continued watching from 30 m away as the veterinary team inspected the scene and the calf. The calf, found to be male, was taking long, deep breaths.
Two and a half hours after the initial observation, the calf was no longer breathing; thus, it was declared dead. Twenty min post-death, F1 returned and touched the calf’s head and body with her trunk. There was no intervention by the forest officials. Using her trunk, she inspected the body for around 1 min, then turned and walked away (ESM_V1d). After moving about 30 m, she turned towards the carcass and stood vigilant as the veterinary team initiated an autopsy (which continued for c. 50 min), after which the carcass was cremated to avoid any potential spread of infection. During cremation (which lasted c. 1 h), F1 circled the area staying around 100 m from the people and the fire. F1’s relationship with the dying calf could not be clearly established, as neither of them was known to us. As the calf was still of suckling age and the female showed no signs of lactation (e.g., no enlarged mammary glands), F1 might not have been the mother. The total duration for peri-mortem observation was 322 min.
Post-mortem observations (reactions to a dead old female)
The behavioural responses of two adult females (F2 and F3: both 30–40 years old) to a recently dead old female were observed on 13 March 2013 at 11.88913°N and 76.20687°E of the BNP. As no elephant carcass was seen at the spot 1 day earlier, the time of death was estimated at less than 14 h earlier, in the late evening of 12 March 2013.
The fresh carcass was spotted near the dried banks of the Kabini reservoir. The research team immediately went to inform the forest officials, who then permitted the team to observe the autopsy to find out the cause of death. When the team returned to the location, two female elephants, F2 and F3, were near the carcass (Fig. 3a). F2 was walking around it in a circular path (two rounds of F2’s movement were witnessed; Fig. 4a–d), while F3 was grazing near the carcass. During her circular movements around the carcass, F2 did not appear to sniff towards the carcass, and her trunk stayed low. She did not attempt to touch the carcass either by trunk or foot (Fig. 4a–d).
So that the autopsy could begin, forest officials chased F2 away from the carcass. As she left, she trumpeted (not recorded), but showed no aggression towards the officials. F3 stayed closer to the carcass (within 30 m) and continued grazing, with no signs of aggression. F2 eventually stopped and stayed near a bamboo thicket, facing towards the carcass and vocalizing occasionally (mostly trumpets, not recorded). When the location around the carcass was explored, it was seen that other elephants had visited the carcass, indicated by the presence of multiple fresh dung boli (no more than 3 h old) of different sizes (circumference 23–49 cm). The presence of some withered ‘green’ leaves and twigs close beside the carcass was also noted (Fig. 3b). The forest officials and a veterinarian conducted the autopsy, during which F3 stayed around 30 m away (Fig. 5). F2 gave sporadic vocalizations from behind a bamboo bush, about 100 m away.
The cause of death was deemed to be a chronic gastrointestinal infection (as the autopsy showed the presence of endoparasites). The carcass was not cremated after the autopsy. The entire observation of the other elephants’ reactions lasted 85 min. After this observation, several scavenger birds and animals were observed feeding on the carcass for a few days. Since the team visited the spot infrequently, there were no sightings of any elephants in the vicinity of the carcass during these periods.
Our observations suggest that Asian elephants perceive distress and respond to dying conspecifics, as reported for their highly social African counterparts and other sentient taxa. The most common thanatological responses that we observed in Asian elephants were epimeletic responses, body-guarding, approach, inspection, visiting, tactile (‘touch’ with trunks and legs), vocalizations (mostly trumpets) and chemosensory behaviours (‘flehmen’ and ‘sniffing’ or ‘directional sniff’/‘sniff-toward’/‘horizontal-sniff’) towards dying and dead conspecifics.
African savannah elephants living in drier habitats such as savanna-woodlands appear to be more gregarious compared to Asian elephants living in dry and moist tropical forest (de Silva and Wittemyer 2012). African savannah elephants live in larger groups with individuals being strongly associated with social units exhibiting a clear hierarchy, whereas Asian elephants live in smaller groups with no clear hierarchical structures (de Silva and Wittemyer 2012; de Silva et al. 2017). Despite differences in their social structures, in general, there seem to be similarities in thanatological responses between these two species. Both species attempt to help and show epimeletic, visitation, approach, inspection, trunk touch, sniffs, vocalizations and body-guarding behaviours towards their injured, dying and dead conspecifics (Moss 1988; Douglas-Hamilton et al. 2006; McComb et al. 2006; Merte et al. 2008; Hawley et al. 2018). While female African elephants have been observed using their tusks to raise or support the dead conspecifics (Douglas-Hamilton et al. 2006), the tuskless female Asian elephants display similar thanatological behaviour without the help of tusks (ESM_V1a).
There also appears to be considerable variability among Asian elephants in response to injured or incapacitated conspecifics. For instance, one of the observed females (Sita, 40–50 years) showed no aggression towards people who approached her injured calf, whereas another female with a dying calf (F1: 15–20 years) behaved agitatedly, vocalized (trumpets and chirps) and charged towards people. Being an old female, Sita may have greater experience in observing the death of conspecifics, and would have interacted with people far more often than F1. Age and previous experiences with dying individuals may, therefore, influence the behavioural responses of Asian elephants to dying conspecifics, as has been documented in several species of non-human primates (see review Watson and Matsuzawa 2018). It is also worth noting that during post-mortem observation, the responding individual did not exhibit any tactile responses (either trunk or feet touches) to a dead female.
Although most thanatological responses recorded in Asian elephants are similar to those of African elephants, we additionally describe vocalizations from a responding individual. Along with behavioural, visual and olfactory signals, vocal cues in particular can play an important role and might reflect the state of a diseased or dying individual (Gonçalves and Carvalho 2019). The vocalizations reported in peri- and post-mortem observations could be signals of an adult’s own distress, as Asian elephants are known to use vocal signals such as chirps, trumpets, roars and rumbles during or in response to distress (Plotnik and de Waal 2014). Although functions of vocal signals towards dying individuals are not clearly understood, there are a few reports describing the types of animal vocalizations and responses associated with it. For instance, vocalizations by responding individuals towards dying individuals can be ‘unique’ calls (Japanese macaques: Green 1975), and could suggest immediate threats or alarm (mainly in corvids, common marmosets, and capuchins; see review Gonçalves and Carvalho 2019), could convey distress (African elephants; Douglas-Hamilton et al. 2006), and sometimes may function as contact or cohesion calls (in ring-tailed lemurs, Sifakas; see review Gonçalves and Carvalho 2019). In the case of peri-mortem observation, the trumpet produced by the responding female F1, in the absence of conspecifics nearby, appeared to be directed towards the calf, to which there was no response from the latter (ESM_V1a). The trumpets produced by F1 towards a dying calf showed a clear decline in duration and varied in intensity. However, the uniqueness of these signals and the reasons for such modulations require further exploration. As the ‘trumpet’ call type is produced in various contexts, such as play or inter-specific aggression in response to the presence of other species (Indian elephants: Nair et al. 2009; Sharma et al. unpublished data) and during disturbance (Sri Lankan elephants: de Silva 2010), the functions of these call types appear to vary widely in Asian elephant society.
The vocalizations observed during the post-mortem observation drew no response from the nearby individuals. Similar observations were reported in ring-tailed lemurs, where other members of the group showed no interest in an injured infant’s calls (Santini 2012). However, McComb et al. (2000) demonstrated that the family of African savannah elephants responded to the playback of their dead female kin even more than two years after death, suggesting a long-term memory associated with their kin. The vocalizations produced by F2 during the post-mortem observation could also be a response directed towards the people who drove her away from the dead female. Nonetheless, these contrasting responses from nearby conspecifics, especially in social animals, could be due to species-specific traits, their genetic relatedness, social structure, association between responding individuals or simply the induced responses.
Besides these behaviours, we also witnessed some seemingly peculiar behaviours, particularly during the post-mortem observation, such as a circular movement (no trunk touches to the carcass) by the responding female, and the presence of withered ‘green’ leaves/twigs and piles of fresh dung boli of different sizes (hinting at visitation by several other individuals) around the carcass. However, being an indirect and single observation, it is difficult to elaborate on the contextual premise of such peculiar behaviour. Further investigations are required to link such behaviours with the elephants’ thanatological responses.
In general, the thanatological responses towards conspecifics in all living elephant species appears to have some striking similarities. Although the divergence time of Loxodonta (African savannah and forest elephants) and Elephas (Asian elephants) is estimated at c. 9–4.2 Mya (Palkopoulou et al. 2018), these similarities could have been ancestral traits retained over the course of proboscidean evolution. Therefore, for better comprehension of thanatological behaviours of Asian elephants, future studies should aim to decode their responses to distress, injuries and death of (1) genetically related and unrelated individuals and (2) heterospecifics in different circumstances [e.g., disease, predation, conflicts with humans (electrocution, poaching or accidents, etc.)]. At the same time, assessing (a) the underlying physiological changes in surviving individuals and (b) vocalizations produced by injured, dying and surviving individuals will further enhance our understanding of the thanatological profile in Asian elephants.
Anderson JR (2016) Comparative thanatology. Curr Biol 26:R553–R556
Anderson JR (2017) Comparative evolutionary thanatology of grief, with special reference to nonhuman primates. Jpn Rev Cult Anthropol 18:173–189
Anderson JR, Biro D, Pettitt P (2018) Evolutionary thanatology. Philos Trans R Soc B 373:20170262. https://doi.org/10.1098/rstb.2017.0262
Appleby R, Smith B, Jones D (2013) Observations of a free-ranging adult female dingo (Canis dingo) and littermates’ responses to the death of a pup. Behav Process 96:42–46
Bates LA, Poole JH, Byrne RW (2008) Elephant cognition. Curr Biol 18:R544–R546
Bearzi G, Eddy L, Piwetz S, Reggente MAL, Cozzi B (2017) Cetacean behavior toward the dead and dying. In: Vonk J, Shackelford TK (eds) Encyclopedia of animal cognition and behavior. Springer, Berlin, pp 1–30. https://doi.org/10.1007/978-3-319-47829-6
Bercovitch FB (2012) Giraffe cow reaction to the death of her newborn calf. Afr J Ecol 51:376–379. https://doi.org/10.1111/aje.12016
Bercovitch FB (2019) A comparative perspective on the evolution of mammalian reactions to dead conspecifics. Primates 20:1–8
de Silva S (2010) Acoustic communication in the Asian elephant, Elephas maximus maximus. Behaviour 1:825–852
de Silva S, Wittemyer G (2012) A comparison of social organization in Asian elephants and African savannah elephants. Int J Primatol 33:1125–1141
de Silva S, Schmid V, Wittemyer G (2017) Fission–fusion processes weaken dominance networks of female Asian elephants in a productive habitat. Behav Ecol 28:243–252
Douglas-Hamilton I, Bhalla S, Wittemyer G, Vollrath F (2006) Behavioural reactions of elephants towards a dying and deceased matriarch. Appl Anim Behav Sci 100:87–102
Gonçalves A, Biro D (2018) Comparative thanatology, an integrative approach: exploring sensory/cognitive aspects of death recognition in vertebrates and invertebrates. Philos Trans R Soc B 16(373):20170263
Gonçalves A, Carvalho S (2019) Death among primates: a critical review of non-human primate interactions towards their dead and dying. Biol Rev Camb Philos Soc 10:15
Green S (1975) Variation of vocal pattern with social situation in the Japanese monkey (Macaca fuscata): a field study. In: Rosenblum LA (ed) Primate behavior, vol 4. Academic, New York, pp 1–102
Hart BL, Hart LA, Pinter-Wollman N (2008) Large brains and cognition: where do elephants fit in? Neurosci Biobehav Rev 32:86–98
Hawley CR, Beirne C, Meyer A, Poulsen JR (2018) Conspecific investigation of a deceased forest elephant (Loxodonta cyclotis). Pachyderm 26:97–100
Iglesias TL, McElreath R, Patricelli G (2012) Western scrub-jay funerals: cacophonous aggregations in response to dead conspecifics. Anim Behav 84:1103–1111
López-Riquelme GO, Fanjul-Moles ML (2013) The funeral ways of social insects. Social strategies for corpse disposal. Trends Entomol 9:71–129
McComb K, Moss C, Sayialel S, Baker L (2000) Unusually extensive networks of vocal recognition in African elephants. Anim Behav 59:1103–1109
McComb K, Baker L, Moss C (2006) African elephants show high levels of interest in the skulls and ivory of their own species. Biol Lett 2:26–28
Merte CE, Gough KF, Schulte BA (2008) Investigation of a fresh African elephant carcass by conspecifics. Pachyderm 46:124–126
Moss CJ (1988) Elephant memories: thirteen years in the life of an elephant family. University of Chicago Press, Chicago
Nair S, Balakrishnan R, Seelamantula CS, Sukumar R (2009) Vocalizations of wild Asian elephants (Elephas maximus): structural classification and social context. J Acoust Soc Am 126:2768–2778
Nandini S, Keerthipriya P, Vidya TN (2017) Seasonal variation in female Asian elephant social structure in Nagarahole-Bandipur, southern India. Anim Behav 134:135–145
Palkopoulou E, Lipson M, Mallick S, Nielsen S, Rohland N et al (2018) A comprehensive genomic history of extinct and living elephants. PNAS 115:E2566–E2574
Plotnik JM, de Waal FB (2014) Asian elephants (Elephas maximus) reassure others in distress. PeerJ 18(2):e278
Plotnik JM, de Waal FB, Moore D III, Reiss D (2010) Self-recognition in the Asian elephant and future directions for cognitive research with elephants in zoological settings. Zoo Biol 29:179–191
Poole JH, Granli P (2011) Signals, gestures, and behavior of African elephants. In: Moss C, Croze H, Lee PC (eds) The Amboseli elephants. University of Chicago Press, Chicago, pp 109–124
Quintana-Rizzo E, Wells RS (2016) Behavior of an adult female bottlenose dolphin (Tursiops truncatus) toward an unrelated dead calf. Aquat Mamm 42:198–202
Santini L (2012) Conflictual behavior in a lemur mother toward a dying infant. Lemur News 16:26–27
Sukumar R (1989) The Asian elephant: ecology and management. Cambridge University Press, New York
Sukumar R (1994) Elephant days and nights. Ten years with the Indian elephant. Oxford University Press, Delhi
Sukumar R (2003) The living elephants: evolutionary ecology, behaviour, and conservation. Oxford University Press, New York
Sukumar R, Suresh HS, Dattaraja HS, John R, Joshi NV (2004) Mudumalai forest dynamics plot, India. In: Losos EC, Leigh EG Jr (eds) Tropical forest diversity and dynamism: findings from a large-scale plot network. University of Chicago Press, Chicago, pp 551–563
Sun Q, Haynes KF, Zhou X (2013) Differential undertaking response of a lower termite to congeneric and conspecific corpses. Sci. Rep 3:1650
Swift K, Marzluff JM (2018) Occurrence and variability of tactile interactions between wild American crows and dead conspecifics. Philos Trans R Soc B 373:20170259
Synchronized Elephant Population Estimation (2017) Statistics from the Project Elephant Division, Ministry of Environment and Forests (MoEF), Government of India
Vidya TN, Prasad D, Ghosh A (2014) Individual identification in Asian elephants. Gajah 40:3–17
Watson CF, Matsuzawa T (2018) Behaviour of nonhuman primate mothers toward their dead infants: uncovering mechanisms. Philos Trans R Soc B 373:20170261
Firstly, we thank all the elephants in our study area for allowing us to observe them peacefully. We would like to thank Mr. Krishna, Mr. Prabhu, Mr. Kethan and Mr. Maadan for helping us to track elephants in the field. We also thank all the forest officials, anti-poaching guards and mahouts at the Bandipur and Mudumalai for providing their valuable support during the field work. We acknowledge the Karnataka and Tamil Nadu Forestry departments for giving us necessary permissions to conduct the research at Bandipur NP and Mudumalai NP, respectively. We also thank Prof. James R. Anderson and Mr. Andre Gonçalves for all the inputs and discussion while preparing this manuscript. We would also like to thank the editor and two anonymous reviewers for providing critical feedback and comments on our manuscript. We would like to thank the Department of Biotechnology (DBT), Government of India, under the DBT-Indian Institute of Science partnership programme, the Rufford Small Grants Foundation, London, UK (RSG13124-1), the Leading Graduate Program in Primatology and Wildlife Science, Kyoto University, Japan, for their financial supports during our fieldwork. RS was a JC Bose National Fellow during the tenure of this work.
Conflict of interest
The authors declare that they have no conflict of interest.
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Electronic supplementary material
Below is the link to the electronic supplementary material.
About this article
Cite this article
Sharma, N., Pokharel, S.S., Kohshima, S. et al. Behavioural responses of free-ranging Asian elephants (Elephas maximus) towards dying and dead conspecifics. Primates 61, 129–138 (2020). https://doi.org/10.1007/s10329-019-00739-8
- Asian elephants
- Dead conspecifics
- Injured calf
- High-frequency vocalizations