Biology:Animal grief

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Grief is a multifaceted response to loss, particularly to the loss of someone that has died, to which a bond or affection was formed.[1] Grief is also witnessed in other animals termed as animal grief, similar to survivor guilt in the human beings. In the late 19th century, Charles Darwin published "The Expression of the Emotions in Man and Animals", in which he has argued that humans and other animals show emotions through remarkably similar behaviours.

Grief was not the focus of animal research until the late 20th and early 21st century. Marc Bekoff, an American biologist has spent his time researching over the emotions in animals, including grief. Studies conducted by Bekoff and other scientists implicate that the following animals could experience grief: Wolves, Chimpanzees,[2] Elephants, Dolphins, Whales,[3] Giraffes,[4] and others.

Animal pain

Pain is a complex experience that involves sensory and emotional components. While physical pain in animals has been well-documented,[5][6][7] it is less clear whether the emotional suffering humans feel can be said to be true of animals.

Animal emotions

Marc Bekoff defines emotion as something that helps behavioral control and management (Bekoff, 2000).[8] It is known that humans have emotions and that it is something fundamental and important in our lives, however it is hard to say if that is true for animals or just some.

Primary emotions and secondary emotions

There are different “categories” of emotions known as primary and secondary emotions. Bekoff explains that primary emotions are similar to reflex or something along the lines of fear or fight-or-flight response. Therefore, it is something that animals react to such as loud noises, unknown objects, or odors (Bekoff, 2000).[8] Having primary emotions is crucial because reactions to these are important for an animal's survival. The part of the brain that is responsible for primary emotions is the limbic system (Bekoff, 2000).[8]

Secondary emotions are part of an experience. These emotions are taking part in the central cortex of the brain because of the requirement of different and higher brain centers. With secondary emotions it allows one (in this case the animals that do feel grief) to create the connection between feelings and actions.

Early research on animal grief

In 1879, Arthur E. Brown studied how a male chimpanzee reacted after the death of his female counterpart. He saw the male chimpanzee display grief and "a cry which the keeper of the animals assures [Brown] he had never heard before").[9] Continuing to the next day, the chimpanzee sulked and barely moved. Brown deciphered that the male chimpanzee was depressed after the female chimpanzee died. However, Brown concluded that any permanent grief is only found in man, as the chimpanzee seemed fine after a couple of days.

William E. Ritter (1925) connects animal and human emotion to provide evidence of human descent from the animal kingdom. He mentions the James-Lange theory, where "all emotional stages as of joy, grief, fear, anger, jealousy, love, are associated with more or less characteristic bodily manifestation".[10] Ritter proposes new evidence to the theory, mentioning that no item on the above list is exclusive to humans, and most are common to the animal world. He argues that because the connection between human emotion and animal emotion is so strong, humans have descended from the animal kingdom.

Marc Bekoff's research

Marc Bekoff is a professor of ecology and evolutionary biology at the University of Colorado-Boulder. He has spent his life studying animals and their emotions, finding that animals grieve quite frequently. Such examples of grieving animals are:

  • Bekoff found that sea lion mothers squeal eerily when they watch their babies being eaten by killer whales.[11]
  • Even after their calves have died, dolphin mothers have been seen to save them and grieve when they confirm the death.[11]
  • Elephants have been "observed to stand guard over a stillborn baby for days with their head and ears hanging down". Orphan elephants, who have watched their mothers be killed, have been shown to wake up screaming.[11] Elephants are known to communally recognize a deceased relative similar to how they greet a newborn, by collectively touching it's corpse or old bones and possibly wailing.
  • A wolf "sniffed [her dead companion], then sat back and gave the most soulful and heart-wrenching howl I've ever heard!"; After a pack member died, the wolves let their tails and heads hang low while walking slowly.[8]
  • Chimpanzee orphans can die while in the state of grieving.[8] Jane Goodall (1990) followed Flint, a chimpanzee, for a few days after the death of Flint's mother Flo. She noted that Flint "walked along one of the branches, then stopped and stood motionless, staring down at an empty nest". Flint had been lethargic, even refusing food. Per Goodall, Flint was "hollow-eyed, gaunt and utterly depressed".[12]
  • Magpies have been seen to mourn over their dead.[13]
  • After a fellow goose died, Konrad Lorenz noted that other geese had their "eyes sink deep into their sockets, and the individual ha[d] an overall drooping experience, literally letting the head hang".[14]

[8] The previous example shows that animals can express their grief and it is possible for an individual to identify that. Bekoff mentions an experience of a bird that lost its partner. It showed many signs of depression before it passed away in a vegetative state similar to the one his partner ended up dying in (Bekoff, 2010). Many animals can show grief when they have lost a loved one of their own.

Duration

One of the questions that can be asked aside from do animals grieve is how long do they grieve for or if these animals show signs of mourning. Anthropologist Barbra J King mentions how animals might sleep less or change their ways in their daily life styles. These animals might also end up staying close to their companion's corpse for a long time as well (Safina, 2015).[15] Something we know is that emotions in a human can change, one day they can be sad and the other happy. Actions are taken days after one of our loved ones dies, such as time off school, work or any social interaction (Safina, 2015).[15] It is actually the same for animals. An example that can be shown is in chimpanzees.

A case is shown with one named Amos. The day before he died he spent the day in his nest and did not move until a female chimpanzee Daisy went up to him (Safina, 2015).[15] When Amos died, one of the chimpanzees who was more sociable than the rest as well as a “higher rank” than the rest did not want to be with the group for weeks. As shown here it also depends on who passed away on how long an animal can grieve. Though through the time spent mourning many signs can appear of an animals grief and agony of the loss of their companion.

Another example mentioned is the dog Hachiko from Tokyo. In this case the dog was not grieving due to the death of an animal, but of the loss of its owner. It is known that this dog went to the station where his owner would come home to every day for around ten years.

Other research

Though Bekoff is the face of recent research on animal grief the field is slowly developing. Some studies have looked into depression in animals, with Paul Willner finding that there are eighteen animal models of depression.[16] Peter J. Fashing & Nga Nguyen (2012) found that a group of chimpanzees groomed and caressed a grieving, older chimpanzee. They also found that "the dying female [elephant] was approached by the matriarch of another group who repeatedly attempted to bring her to her feet using her tusks". Even otters grieve, where members of a group caught fish for months and brought them to the matriarch who had failing vision and poor mobility.[17]

Animal mourning

In social groups, bereavement is a natural stress reaction in the event of the loss of a significant loved one through death.[18] Like Humans, social animal species are affected by the loss of one of their own and can undergo psychological stress or trauma.[19]

The behaviour that these social groups expect following bereavement is referred to as mourning: the act of sorrowing during a period of time during which signs of grief are shown.[18][20]

Ample research in fields of biology, animal behaviour, evolutionary and ecological biology, cognitive ethology, and neuroscience has shown that animals have brains and minds, used, like in humans, for thinking and feeling.[21] As such they show moods and emotions and therefore grief and empathy.[21] However, our understanding of the putative emotional responses to loss of animals is limited by communication capabilities and possible differences between our psychological responses to death and theirs.[22] Thanatology used to refer to the "academic and often scientific study of death in human beings". It has now been broadened to include the study of the mechanism, body changes, but also the psychological and social consequences of death among other species.[19]

Social emotions shared in mammals

Primitive tendencies to nurturance probably evolved before the divergence of mammalian and avian stock from their common ancestor. For instance, recent paleontological evidence suggests that some dinosaurs may have also exhibited maternal care.[21] However maternal devotion, and complex social feelings in mammals are sure to have emerged with the evolution of the limbic system.[21] One of the greatest advances in the evolution of emotionality was the ability of the young to value social support. Jaak Panksepp says that this is achieved by a separation-distress or PANIC system, which is a sensitive emotional barometer that keeps track of the level of social support they are receiving. When social contact is lost, organisms experience the painful feeling of separation, and the young reacts vigorously in an attempt to reestablish contact and care.[23][21]

Scientists documenting social animals nurturant behaviour towards dead young explain it by the species gregarious nature.[24][25] These species strongly rely on cooperation and social bonding, including allomothering,[26][25] explaining such actions as adults taking care of other parent's calves and even adoption.[27][25] Human infants are born with brains that are about 23 percent of their final capacity, characteristic that neuroanatomists explain by the yet need to learn.[21] This case is similar in some species, such as in elephants[21] or odontocetes,[28][27] and these exhibit long periods of infant dependency and thus substantial nurturant behaviours in mothers.[27] Fred B. Bercovitch states that among female mammals, lifetime reproductive success depends upon rearing, more than on production of offsprings,[29] which is associated with the strong maternal-infant bond.[29] He then proposes that oxytocin's role in this mother-infant bond might have evolved to be implicated in promotion of social bonds as well. This intricate connection between physiology and behaviour may allow the rationale for the care and carrying of deceased newborns found behaviour, in group-living species.[19][29]

Mourning animals

Elephants

Elephants are a highly cognitively and emotionally engaged animal.[21][30] “Through observational evidence they seem to have a real unusual interest in the dead of their own species either fresh carcasses or skulls. [...] and the interest seems to persist long after death” - says Karen McComb, co-director of the Mammal Vocal Communication and Cognition Research Group at the University of Sussex. She is a behavioural ecologist and an expert in analyzing vocalizations of mammals to assessing thoughts and emotions.[21][31] For that, McComb studies 'contact calls', powerful rumbles elephant use to keep in contact for large distances, and plays recordings of these to approaching herds of elephants while observing their reaction to them. These are her clues in her study. When played the call of a missing family member, the elephants were seen running trumpeting towards the speaker, as if they had someone in ‘mind’ they expected to see.[21] She also used the call of a fifteen-year-old female elephant who had died, and played it twice to her family, once a few months after her death and then twenty-three month later. Similarly, the herd ran to her rumbling in greeting, walking directly to the speaker. “They hadn't forgotten her. [...] But I was uneasy doing that test.” - she mentions.[21] Elephants have intensive social natures and retain their memories for many years.[32] They have the adequate neural anatomy for long-term memory: their brains have large and complex frontal lobes, important structures for storing and retrieving memories of scent, touch, smell, and sound.[21] Poachers often target the matriarchs or older females of the group because they have larger tusks. This is an impactful loss for the group because they lose a lifetime of learning and knowledge.[21]

Elephants have a concept of death. For instance, they recognize carcasses and skeletons of their own kind as well as of others.[33][21] McComb performed experiments in which she placed skulls of elephants killed by poacher, to make miniature elephant graveyards, in the path of approaching herds. She observed that they took a real interest in skulls and bones of their own kind, touching and investigating the body for hours.[31] The behaviour they adopt when encountering their dead relatives varies, says Cynthia Moss, ethologist and director of the Amboseli Elephant Research Project, and who invited McCom to join her team. “They stop and become quiet and tense in a different way from anything I have seen in other situations”.[21] Responses of elephants to conspecific carcasses at various stages post-mortem has been intensively documented.[32][34] They display touching behaviours such as smelling and caressing the remaining body with their hind feet or with their trunk. Initially, the dead might be treated as if it was still alive, or unable to move as if injured. It gets carried in order to keep up with the group, tried to be lift up and help stand. Such actions reveal the elephant's empathetic and compassionate nature [21] and shows that they do not want to abandon the deceased individual.[29] They have been seen to cover bodies with nearby vegetation, as a burial.[32] Out of many others, Sharma [35] described a case of an adult female mother in southern India, standing around her dead calf for weeks. The mother later watched from a distance a veterinary team conducting an autopsy on-site and remained around 100m away from the fire on which the calf's body was cremated.[32][35]

There are tales about elephant's graveyards.[32][21] These are believed to be places where dying elephants direct themselves to die with their conspecifics. Today it is believed that these are actually the remainder bones and skeletons of elephants killed by poachers. However, some reports allow the hypothesis that elephants, which are always on the move, may return occasionally to places where a member has been buried.

Cetaceans

Many studies reviewed reports of whales and odontocetes, dolphins in particular, displaying nurturant behaviours towards dead calves and juveniles, either by supporting or carrying the young at the surface for very long distances and for a very long time.[25]

One example of the many cases Reggente [25] reports in their study describes an adult Risso's dolphin interacting with a dead calf. Observers related the animal swimming holding on its back the young dead by its dorsal fin, then swimming in circles under the carcass, touching it with several parts of its body (rostrum, and pectoral and dorsal fin), and remaining in vertical position next to the dead calf.[25] Later, the adult pushed the body away from its observers, who did not attempt to follow them. Another reports an adult Indo-Pacific bottlenose dolphin pushing a small dead dolphin across waters. Biologists decided to fix a rope and drag the carcass to the shore to bury it. Observers report having the adult following it, touching it and swimming around until it reached shallow waters.[25] In one study, Kilborn (1994)[36] reported a case in Point Defiance Zoo and Aquarium's whale exhibit, in which a captive mother beluga whale, Mauyak, gave birth to a male calf which died right after birth. After delivering the placenta a few hours later, she took it in her mouth and moved out of reach from the staff, carrying it for hours before it was removed from the pool. The day after, she was found carrying a 30-cm diameter pink buoy that had been available to her prior to the birth.[36] Many testimonies also report seeing killer whales carrying dead calves for miles.

Smith and Sleno (1986),[37] and Kilborn (1994)[36] allow the hypothesis that these recurrent nurturant behaviours found in the wild might be due from the incapacity of the mothers to accept their infants' death.

Non-human primates

Specialist Jane Goodall recognizes that chimpanzees have distinct personalities that impact their success in life.[21] As the animals discussed previously and still many others, they show sophisticated behaviours: they forge friendship, communicate, have a sense of fairness, occasionally deceive and lie, and grieve for their dead.[31]

Concerning great apes, the best known reports until today probably account Goodall's descriptions of chimpanzee mothers carrying and caring for dead infants,[38] as well as reports from Geza Teleki on a group's response to an accidental adult chimpanzee's fatal fall from a tree.[39][19] Anderson et al.[40] report was the first to include video evidence of chimpanzees' precise moment response at the death of an elderly female of their group.[19] They could draw parallels with behaviours seen in humans in such circumstances, such as pre-death care of the female, close inspection and testing for signs of life at the moment of death, male aggression towards the corpse, all-night attendance by the deceased's adult daughter, cleaning the corpse, and later avoidance of the place where the death occurred.[40]

In humans, expressions of sympathy and moral support may arise towards the family members in the event of a relative's death.[19] Goldsborough [41] observed captive chimpanzees' affiliative interactions with an adult female before and after the birth of her stillborn baby. She received more affiliation than before, including from previously non-affiliative chimpanzees, and particularly from a female who also had a stillborn infant in the years before her.[15]

Giraffes

In Giraffes, the mother-infant bond that happens during the neonatal period lasts for a minimum of 12–16 months [42] and there are suggestions that mother and daughters may associate for several years and more.[43] Giraffe's calves have high mortality rate (58-73%)[44] in the first year and literature shows mothers adopt a characteristic nurturant behaviour in the event of their calf's death. Cases have described behaviours of mothers standing guard over their newborn dead calves,[29] while giraffes rarely remain stationary for long,[29] or of mothers watching quietly from distance their infant carcasses being fed on by predators.[32]

Laysan albatrosses

Laysan albatrosses are monogamous. They don't breed until they are eight or nine years old. John Klavitter is a U.S fish and Wildlife Service biologist at Midway and says that when an albatross loses one of their mate the surviving bird “goes through a year or two of a mourning period” before it starts searching again for a new mate, a process that can take many years.[21]

Mute swans

Mute swans, Cygnus olor, are monogamous, and are known to go through a mourning period[45] following the death or loss of their mate or a cygnet.[46] Should a parent die while there are cygnets present, the remaining adult will take up its counterpart's duties in caring for the offspring. After undertaking the mourning period, the surviving swan, if it is alone, may remain in the area or depart to locate a nearby flock. They have been noted to develop emotional attachments to their mates and other individuals.[47]

Other animals

In human management systems, such as in parks, farms and laboratory animals, animals are subject to abrupt separation from their mothers[48] and other conspecifics.[49][22] McGraph [22] addresses the abundant scientific findings about grief and separation distress that are found in domesticated animals such as cows, pigs, sheep, goats, horses and chickens. Physiological changes resulting from it include distress vocalizations, increased activity or locomotion, escape attempts, increased heart-rate and increased hypothalamic-pituitary activity.[22] They do not show sign of the bi-phasic protest-despair reaction typical of grief.[22] Nonetheless, this does not mean that they are not capable of grieving as some may present other reactions. This assumption raises issues and debates about ethics and welfare of animals in situations such as with laboratory animal management.

William Russel and Rex Burch published the seminal book, The Principles of Humane Experimental Techniques,[50] and introduced the now referred "3Rs" principles: reduction, refinement and replacement of animal use. Despite the attention brought to this issue by the authors and since, the number of animals used in research and testing continues to increase drastically.[51]

Marc Bekoff is a former Professor of Ecology and Evolutionary Biology at the University of Colorado, and co-founder with Jane Goodall of Ethologists for the Ethical Treatment of Animals.[52] He supports that animals have emotions, empathy and moral intelligence, a theory that has been extensively supported by scientific proofs.[53] In his ample work, some of his reports include cases of grieving animals. Among others: magpies standing next to a conspecific's corpse, gently pecking at it, seems to be a ritual that has been observed in ravens and crows as well; baboons; elephants.[53]

Human grief from pet-loss

Some examples of empathy cross the species barrier. One is the famous dog-human relationship.

The process of grief over the death of a pet among owners is under-explored in research and often devalued, but some studies have tackled on comprehensive analysis of grief responses in dog owners, showing that the more the owner has the tendency to humanize his animal the more intense the grief experienced will be.[18] In consequence, the mourning behaviour will be similar to ones that he would have if he went through the loss of a human companion: burials, cremations, taxidermies. These mourning rituals similarly allow owners to recapture and re-experience intensely lived moments of comradeship and closeness with their fellow companion.[54] When the pet may have played an additional role such as being a working dog, service or therapy animal, owners will not only grieve the loss of a companion but also the loss of a co-worker.

Pet cemeteries exist. One is the Psi Los (“Dog's fate”) cemetery located in Konik Nowy, on the outskirts of the Polish capital city of Warsaw. There, the place is home to thousands of animal graves and tombstones, including numerous species: dogs, but also cats, rats, rabbits, guinea pigs, hamsters, mice, ferrets, chinchillas, canary birds, turtles, and at least one iguana. In social practices, significant similarities can be observed in ways to commemorate companion animals to those typically found in human cemeteries in Poland: families paying a visit, lighting candles, placing fresh or artificial flowers. Moreover, at Psi Los, the Catholic All Saint's celebration is mirrored on the same day (November 1) by the informal Pet Memorial day.[55]

Implications of research

Though grief in animals may seem questionable, evidence shows it is abundant. From chimpanzees to otters to sea lions, animals grieve just like humans do. Researchers like Bekoff, Fashing, Nguyen, and others, are studying every day to help better understand how and why animals grieve. With increased knowledge, humans can have better relationships with animals. For example, zoo caretakers can study chimpanzee grief habits and better notice when a chimpanzee mother is mourning. The caretakers can then help the mother cope and live a healthy and successful life. Research shows grief in animals, and understanding that can help humans form closer, healthier connections with them.

References

  1. ↑ McMaryion, Anthony (2020-12-15) (in en). How Long the Night. Christian Faith Publishing, Inc.. ISBN 978-1-0980-5340-6. https://books.google.com/books?id=oJMuEAAAQBAJ&dq=a%20multifaceted%20response%20to%20loss,%20particularly%20to%20the%20loss%20of%20someone%20or%20something%20that%20has%20died,%20to%20which%20a%20bond%20or%20affection%20was%20formed&pg=PT20. 
  2. ↑ "Chimps Understand and Mourn Death, Research Suggests" (in en). 2010-04-26. https://www.livescience.com/6335-chimps-understand-mourn-death-research-suggests.html. 
  3. ↑ Magazine, Smithsonian; Daley, Jason. "Study Suggests Dolphins and Some Whales Grieve Their Dead" (in en). https://www.smithsonianmag.com/smart-news/study-suggests-dolphins-and-some-whales-grieve-their-dead-180969414/. 
  4. ↑ Magazine, Smithsonian; Nuwer, Rachel. "Is This Mother Giraffe Mourning Her Dead Baby?" (in en). https://www.smithsonianmag.com/smart-news/is-this-mother-giraffe-mourning-her-dead-baby-29645135/. 
  5. ↑ "The Surprisingly Humanlike Ways Animals Feel Pain" (in en). 2016-12-03. https://www.nationalgeographic.com/animals/article/animals-science-medical-pain. 
  6. ↑ Lopez, German (2017-01-23). "Animals can feel pain. A biologist explains how we know." (in en). https://www.vox.com/science-and-health/2017/1/23/14325172/animals-feel-pain-biologist. 
  7. ↑ Animals, National Research Council (US) Committee on Recognition and Alleviation of Pain in Laboratory (2009) (in en). Pain in Research Animals: General Principles and Considerations. National Academies Press (US). https://www.ncbi.nlm.nih.gov/books/NBK32655/. 
  8. ↑ 8.0 8.1 8.2 8.3 8.4 8.5 Bekoff, Marc (2010) (in en). The Emotional Lives of Animals: A Leading Scientist Explores Animal Joy, Sorrow, and Empathy--and why They Matter. New World Library. ISBN 9781577313489. https://books.google.com/books?id=TIW2D7eR_Y8C. 
  9. ↑ Brown, Arthur E. (1879). "Grief in the Chimpanzee". The American Naturalist 13 (3): 173–175. doi:10.1086/272298. 
  10. ↑ Ritter, William E. (1925). "The Emotions in Man and Animals". The Scientific Monthly 21 (2): 137–138. Bibcode: 1925SciMo..21..137R. 
  11. ↑ 11.0 11.1 11.2 Bekoff, Marc (2000). "Animal Emotions: Exploring Passionate Natures | BioScience | Oxford Academic" (in en). BioScience 50 (10): 861. doi:10.1641/0006-3568(2000)050[0861:AEEPN2.0.CO;2]. 
  12. ↑ Goodall, Jane (2011) (in en). Through a Window: 30 years observing the Gombe chimpanzees. Orion. ISBN 9780297865360. https://books.google.com/books?id=Gc59YpamCvgC. 
  13. ↑ Bekoff, Marc (2009). "Animal emotions, wild justice and why they matter: Grieving magpies, a pissy baboon, and empathic elephants - ScienceDirect" (in en). Emotion, Space and Society 2 (2): 82–85. doi:10.1016/j.emospa.2009.08.001. 
  14. ↑ Bekoff, Marc (2002) (in en). Minding Animals: Awareness, Emotions, and Heart. Oxford University Press, USA. ISBN 9780195150773. https://books.google.com/books?id=Or4TDAAAQBAJ. 
  15. ↑ 15.0 15.1 15.2 15.3 "The Depths of Animal Grief — NOVA Next | PBS" (in en-US). NOVA Next. 2015-07-08. http://www.pbs.org/wgbh/nova/next/nature/animal-grief/. 
  16. ↑ Willner, Paul (1984). "The validity of animal models of depression". Psychopharmacology 83 (1): 1–16. doi:10.1007/BF00427414. PMID 6429692. 
  17. ↑ "Behavior toward the dying, diseased, or disabled among animals and its relevance to paleopathology" (in en). http://anthro.fullerton.edu/_resources/pdf/nga/FashingandNguyen2011.pdf. 
  18. ↑ 18.0 18.1 18.2 Uccheddu; De Cataldo; Albertini; Coren; Da Graça Pereira; Haverbeke; Mills; Pierantoni et al. (2019-11-07). "Pet Humanisation and Related Grief: Development and Validation of a Structured Questionnaire Instrument to Evaluate Grief in People Who Have Lost a Companion Dog" (in en). Animals 9 (11): 933. doi:10.3390/ani9110933. ISSN 2076-2615. PMID 31703410. 
  19. ↑ 19.0 19.1 19.2 19.3 19.4 19.5 Anderson, James R. (January 2020). "Responses to death and dying: primates and other mammals" (in en). Primates 61 (1): 1–7. doi:10.1007/s10329-019-00786-1. ISSN 0032-8332. PMID 31902088. 
  20. ↑ "Mourning". https://www.merriam-webster.com/dictionary/mourning. 
  21. ↑ 21.00 21.01 21.02 21.03 21.04 21.05 21.06 21.07 21.08 21.09 21.10 21.11 21.12 21.13 21.14 21.15 21.16 21.17 21.18 Morell, Virginia (2013). Animal wise how we know animals think and feel. Broadway Books. ISBN 978-0-307-46145-2. OCLC 1028468220. 
  22. ↑ 22.0 22.1 22.2 22.3 22.4 McGrath, N; Walker, J; Nilsson, D; Phillips, C (2013-02-01). "Public attitudes towards grief in animals". Animal Welfare 22 (1): 33–47. doi:10.7120/09627286.22.1.033. ISSN 0962-7286. 
  23. ↑ Panksepp, Jaak (2005). Affective neuroscience : the foundations of human and animal emotions. Oxford University Press. ISBN 978-0-19-517805-0. OCLC 793022084. 
  24. ↑ Mann, Janet. (2000). Cetacean societies : field studies of dolphins and whales. University of Chicago Press. ISBN 0-226-50340-2. OCLC 42309843. 
  25. ↑ 25.0 25.1 25.2 25.3 25.4 25.5 25.6 Reggente, Melissa A. L.; Alves, Filipe; Nicolau, Cátia; Freitas, Luís; Cagnazzi, Daniele; Baird, Robin W.; Galli, Paolo (2016-05-24). "Nurturant behavior toward dead conspecifics in free-ranging mammals: new records for odontocetes and a general review". Journal of Mammalogy 97 (5): 1428–1434. doi:10.1093/jmammal/gyw089. ISSN 0022-2372. 
  26. ↑ Bates, Lucy A.; Lee, Phyllis C.; Njiraini, Norah; Poole, Joyce H.; Sayialel, Katito; Moss, Cynthia J.; Byrne, Richard W. (2008). "Do Elephants Show Empathy?" (in en). Journal of Consciousness Studies 15 (10–11): 204–225. ISSN 1355-8250. http://dspace.stir.ac.uk/handle/1893/946. 
  27. ↑ 27.0 27.1 27.2 MANN, JANET; SMUTS, BARBARA B. (May 1998). "Natal attraction: allomaternal care and mother–infant separations in wild bottlenose dolphins". Animal Behaviour 55 (5): 1097–1113. doi:10.1006/anbe.1997.0637. ISSN 0003-3472. PMID 9632497. 
  28. ↑ Brodie, Paul F. (July 1969). "Duration of Lactation in Cetacea: An Indicator of Required Learning?". American Midland Naturalist 82 (1): 312–314. doi:10.2307/2423849. ISSN 0003-0031. 
  29. ↑ 29.0 29.1 29.2 29.3 29.4 29.5 Bercovitch, Fred B. (2019-03-20). "A comparative perspective on the evolution of mammalian reactions to dead conspecifics". Primates 61 (1): 21–28. doi:10.1007/s10329-019-00722-3. ISSN 0032-8332. PMID 30895413. 
  30. ↑ Wylie, Dan. (2018). Death and compassion : the elephant in Southern African literature. Wits University Press. ISBN 978-1-77614-220-0. OCLC 1109783548. 
  31. ↑ 31.0 31.1 31.2 Packham, Chris. "Inside the Animal's Mind: Secrets of the Social World". https://www.bbc.co.uk/programmes/b03vd19b. 
  32. ↑ 32.0 32.1 32.2 32.3 32.4 32.5 Anderson, James R. (January 2020). "Responses to death and dying: primates and other mammals". Primates 61 (1): 1–7. doi:10.1007/s10329-019-00786-1. ISSN 0032-8332. PMID 31902088. 
  33. ↑ McComb, Karen; Baker, Lucy; Moss, Cynthia (2005-10-25). "African elephants show high levels of interest in the skulls and ivory of their own species". Biology Letters 2 (1): 26–28. doi:10.1098/rsbl.2005.0400. ISSN 1744-9561. PMID 17148317. 
  34. ↑ Goldenberg, Shifra Z.; Wittemyer, George (2019-11-11). "Elephant behavior toward the dead: A review and insights from field observations". Primates 61 (1): 119–128. doi:10.1007/s10329-019-00766-5. ISSN 0032-8332. PMID 31713106. 
  35. ↑ 35.0 35.1 Sharma, Nachiketha; Pokharel, Sanjeeta Sharma; Kohshima, Shiro; Sukumar, Raman (2019-08-19). "Behavioural responses of free-ranging Asian elephants (Elephas maximus) towards dying and dead conspecifics". Primates 61 (1): 129–138. doi:10.1007/s10329-019-00739-8. ISSN 0032-8332. PMID 31428950. 
  36. ↑ 36.0 36.1 36.2 Kilborn, Sherrie S. (October 1994). "Object Carrying in a Captive Beluga Whale (Delphinapterus Leucas) as Possible Surrogate Behavior". Marine Mammal Science 10 (4): 496–501. doi:10.1111/j.1748-7692.1994.tb00510.x. ISSN 0824-0469. 
  37. ↑ Smith, T. G.; Sleno, G. A. (1986-07-01). "Do white whales, Delphinapterus leucas, carry surrogates in response to early loss of their young?". Canadian Journal of Zoology 64 (7): 1581–1582. doi:10.1139/z86-237. ISSN 0008-4301. 
  38. ↑ Van Lawick-Goodall, Jane (1968). "The Behaviour of Free Living Chimpanzees in the Gombe Stream Reserve". Animal Behaviour Monographs 1, part 3: 161–311. 
  39. ↑ Teleki, Geza (1973). "Group Response to the Accidental Death of a Chimpanzee in Gombe National Park, Tanzania". Folia Primatologica 20 (2–3): 81–94. doi:10.1159/000155569. ISSN 0015-5713. PMID 4789109. 
  40. ↑ 40.0 40.1 Anderson, James R.; Gillies, Alasdair; Lock, Louise C. (April 2010). "Pan thanatology". Current Biology 20 (8): R349–R351. doi:10.1016/j.cub.2010.02.010. ISSN 0960-9822. PMID 21749950. 
  41. ↑ Goldsborough, Zoë; van Leeuwen, Edwin J. C.; Kolff, Kayla W. T.; de Waal, Frans B. M.; Webb, Christine E. (2019-09-04). "Do chimpanzees (Pan troglodytes) console a bereaved mother?". Primates 61 (1): 93–102. doi:10.1007/s10329-019-00752-x. ISSN 0032-8332. PMID 31485897. 
  42. ↑ Strauss, Megan K. L.; Muller, Zoe (2012-11-16). "Giraffe mothers in East Africa linger for days near the remains of their dead calves". African Journal of Ecology 51 (3): 506–509. doi:10.1111/aje.12040. ISSN 0141-6707. 
  43. ↑ Pratt, David M.; Anderson, Virginia H. (2010-04-26). "Giraffe Cow-Calf Relationships and Social Development of the Calf in the Serengeti". Zeitschrift für Tierpsychologie 51 (3): 233–251. doi:10.1111/j.1439-0310.1979.tb00686.x. ISSN 0044-3573. 
  44. ↑ FOSTER, J. B.; DAGG, A. I. (May 1972). "Notes on the biology of the giraffe". African Journal of Ecology 10 (1): 1–16. doi:10.1111/j.1365-2028.1972.tb00855.x. ISSN 0141-6707. 
  45. ↑ "The mute swan | Swan Lifeline" (in en-GB). https://www.swanlifeline.org.uk/the-mute-swan/. 
  46. ↑ "The care and treatment of swans and waterfowl with an established worldwide reputation." (in en-GB). https://www.theswansanctuary.org.uk/general-information/. 
  47. ↑ "Mute Swan - NYS Dept. of Environmental Conservation". https://www.dec.ny.gov/animals/7076.html. 
  48. ↑ Newberry, R.; Swanson, J. (2001), "Breaking social bonds.", Social behaviour in farm animals, CABI, pp. 307–331, doi:10.1079/9780851993973.0307, ISBN 978-0-85199-397-3 
  49. ↑ Siebert, Katrin; Langbein, Jan; Schön, Peter-Christian; Tuchscherer, Armin; Puppe, Birger (April 2011). "Degree of social isolation affects behavioural and vocal response patterns in dwarf goats (Capra hircus)". Applied Animal Behaviour Science 131 (1–2): 53–62. doi:10.1016/j.applanim.2011.01.003. ISSN 0168-1591. 
  50. ↑ William M. S. Russel, Rex Leonard Burch (1959). The Principles of Humane Experimental Techniques. London: Methuen. 
  51. ↑ Ferdowsian, Hope R.; Beck, Nancy (2011-09-07). "Ethical and Scientific Considerations Regarding Animal Testing and Research". PLOS ONE 6 (9): e24059. doi:10.1371/journal.pone.0024059. ISSN 1932-6203. PMID 21915280. Bibcode: 2011PLoSO...624059F. 
  52. ↑ "ANNOUNCEMENT" (in en). Journal of Applied Animal Welfare Science 3 (3): 277. July 2000. doi:10.1207/S15327604JAWS0303_11. ISSN 1088-8705. 
  53. ↑ 53.0 53.1 Bekoff, Marc (December 2009). "Animal emotions, wild justice and why they matter: Grieving magpies, a pissy baboon, and empathic elephants". Emotion, Space and Society 2 (2): 82–85. doi:10.1016/j.emospa.2009.08.001. ISSN 1755-4586. 
  54. ↑ Courtney, Julia (2018-12-01). "Animal Objects: Memory, Desire and Mourning". Cahiers victoriens et édouardiens (88 Automne). doi:10.4000/cve.4192. ISSN 0220-5610. 
  55. ↑ DeMello, Margo (August 2016). Mourning animals : rituals and practices surrounding animal death. ISBN 978-1-61186-212-6. OCLC 930364167. 




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