Introduction

Higanbana flower (Lycoris radiata)“A few days is all it takes for thisJapanese Crimson beauty to make its
appearance from the Summer shadows.
All parts of the plant are toxic tohumans and other animals. The bulbs
were historically planted on top ofburial sites to discourage animals fromdisturbing the body which would havebeen laid directly in the earth“
Adam Martin (2022)

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Welcome to the 2024 edition of the International Symposium on Comparative Evolutionary
Thanatology (EvoThan) hosted by the EvoThan Collective. We are thrilled to announce our
event, taking place in the serene setting of Kyoto, Japan, from Thursday, June 20th, to
Friday, June 21st, 2024. Comparative thanatology is the interdisciplinary study of deathrelated behaviors and processes across different animal species, aiming to elucidate
evolutionary, ecological, and social factors influencing animals' responses towards mortality.
The EvoThan Symposium will be a gathering of experts, researchers, and enthusiasts
dedicated to exploring and advancing our understanding of such animal reactions to deathand dying.
The EvoThan Collective is an international non-profit organization dedicated to advancing
the scientific understanding of comparative evolutionary thanatology—the study of deathrelated behaviors across human and nonhuman animals. Welcoming researchers fromdiverse fields, including biology, ethology, anthropology, and comparative psychology, the
EvoThan Collective emphasizes rigorous scientific methods to explore the nature andevolution of death-related phenomena. Through conferences, publications, and acommitment to the scientific method, it serves as a collaborative platform forinterdisciplinary knowledge exchange in the evolving field of comparative evolutionary
thanatology.
Recognizing the diverse needs of our global community, the EvoThan Symposium will be a
hybrid event, ensuring accessibility for participants who are unable to join us in person. Join
us to engage with eminent scholars and researchers involved in comparative thanatology, as
they share groundbreaking insights and discoveries. Whether you're presenting your own
research or participating in lively discussions, your contributions will shape the dialogue of
this international gathering.

EvoThan Collective.
June, 2024

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Speakers“The {anthill] passage leading downward from theopening is not all straight or easy for any othercreature to traverse; it passes through turns andtwists with branching tunnels and connecting
galleries and terminates in three hollow cavities. Oneof these is their common dwelling-place, another astoreroom for provisions, while in the third they
deposit the dying.“
Plutarch (46-74 CE)

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Susan Carey

Barbara J. King

Emerita Professor Susan Carey’s work at Harvard University
has focused on the development of concepts in children andadults. Her research explores the initial representational
repertoire, developmental changes, and learning mechanisms
underlying shifts in conceptual understanding. Of particularinterest are her groundbreaking studies on children's
acquisition of the concept of death. Recognized withnumerous awards, Prof. Carey has had a significant impact in
the fields of developmental psychology and cognition.

Emerita Professor Barbara J. King of William & Mary’s
Anthropology Department is now a full-time science writerand public speaker. She has written 7 books on animals’ lives,
including How Animals Grieve, and numerous magazine
essays. Her TED talk on love and grief in the animal kingdom
has been viewed over 3.5 million times. Prof. Barbara King's
website is www.barbarajking.com

James R. Anderson

Alecia Carter

Emeritus Professor James Anderson at Kyoto University, isinternationally recognized for his research on wild and captive
primates, particularly in the realms of primate cognition andmirror self-recognition. His substantial contributions to thefield endure, highlighted by his editorial roles in esteemedjournals. Notably, he has edited two special journal editions
on comparative thanatology and thrice-organized the Kyoto
Workshop on Evolutionary Thanatology, showcasing his
authority in the research of primate cognition and its
implications for the understanding of comparative
evolutionary thanatology.

Prof. Alecia Carter is a Lecturer in Evolutionary
Anthropology at the Department of Anthropology,
University College London, UK, focuses her research on how
primates, particularly baboons, respond to the deaths ofothers, offering insights into the evolution of cognition andemotion. Her extensive study at the Tsaobis Baboon Projectin Namibia has revealed intriguing behaviors, such as baboon
mothers carrying and grooming the corpses of their infants,
shedding light on the evolutionary origins of grief and deathawareness.

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Takahisa Miyatake

Teresa Iglesias

Prof. Takahisa Miyatake, a distinguished researcher at
Okayama University, is a leading expert in insect behavior
with over 170 publications. With years of dedicated andextensive research on the evolutionary aspects of deathfeigning behavior in insects, his work provides valuable
insights into the emergence of anti-predator strategies,
emphasizing the trade-offs individuals face between
enhancing survival prospects and limiting opportunities forfeeding and reproduction.

Dr. Teresa Iglesias serves as the team leader for cephalopod
support at the Okinawa Institute of Science and Technology.
Her research centers on behavior, ecology, and neuroscience,with a specific emphasis on the investigation of sleep behavior
in cephalopods. Previously, Dr. Iglesias conducted anexperimental study on thanatological responses in western
scrub jays, which raised intriguing questions about the socialand emotional dimensions of these mobbing interactions,shedding light on the complex behaviors exhibited by scrub
jays when faced with both dead conspecifics andheterospecifics

Antonio Osuna-Mascaro´
Dr. Antonio Osuna-Mascaró is an animal cognition
researcher with a diverse academic background that include
paleontology and tool use in wild chimpanzees. Currently
based at the Goffin Lab, Messerli Research Institute,
Vetmeduni, his work focuses on unraveling the cognitive andtool-using abilities of Goffin's cockatoos. Among these is his
latest research to assess their understanding of irreversibility inrelation to the concept of death.

André Gonçalves
Dr. André Gonçalves, a post-doctoral researcher at the Centerfor the Evolutionary Origins of Human Behavior (formerly
Primate Research Institute) at Kyoto University, is deeply
engaged in investigating the comparative thanatology ofprimates and other non-human animals through experimental
methods. His research primarily focuses on exploring how
chimpanzees perceive and process death and death-related
stimuli. Utilizing techniques such as touchscreen tasks andeye-tracking technology, he aims to uncover the intricacies ofchimpanzee cognition in this domain.
.

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Nachiketha Sharma
Dr. Nachiketha Sharma, a postdoctoral fellow at the Kyoto University Institute forAdvanced Study, is deeply interested in understanding Asian elephants from a holistic
perspective. His research mainly focuses on ecology, acoustic communication,
physical cognition, and thanatology of Asian elephants both in wild and captive
conditions.

Sanjeeta Sharma Pokharel
Dr. Sanjeeta Sharma Pokharel is an elephant biologist who has
been exploring the physiological adaptive mechanisms
concerning ecological and human-induced challenges in freeranging and captive Asian elephants. Her primary researchinterests revolve around understanding physiological andbehavioural interplays in animals. A part of her elephantresearch also focuses on comparative thanatological studies in
Asian elephants. She is a member of the IUCN Asian
Elephant Specialist Group and currently works as a JSPS
postdoctoral fellow at Asian and African Area Studies(ASAFAS), Kyoto University.

Leanne Proops
Dr. Leanne Proops is an Associate Professor in the Centre for Comparativeand Evolutionary Psychology, University of Portsmouth, UK. She studies
the socio-cognitive abilities of domestic equids (horses, donkeys and mules)
and is fascinated by the field of evolutionary thanatology. Her current
projects include an exploration of domestic horse and feral donkey responses
to the carcasses and bones of their species..

Claudia Stephan

Qian Sun
Dr. Qian “Karen” Sun is an Assistant Professor in the Department of Entomology atLouisiana State University. Her research centers on the chemical and behavioral
ecology of social insects, with a primary emphasis on termites. Her earlier
investigations have delved into the behavioral responses of termites to deadconspecifics and the mechanisms of death recognition. Additionally, she has
contributed to the field with two review papers on corpse management in social
insects. She remains interested in the intriguing research topic of evolutionary
thanatology.

Dr. Claudia Stephan is a lecturer and researcher at the
Department of Biology at the Friedrich-Alexander University
Erlangen-Nürnberg, Germany. Her research primarily
focuses on the evolution and the development of vocal
communication and cognition in forest guenons. Dr.
Stephan recently added topics on the behaviour andunderlying cognitive capacities in forest elephants and gorillasin their natural habitats to her research portfolio. She isparticularly interested in how highly social species respond to
deceased conspecifics and what kind of (contextual)
information is inferred during such encounters.

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Abstracts:
Main Talks“After the death of the female, which took place
early in the morning, the remaining one mademany attempts to rouse her, and when he foundthis to be impossible his rage and grief were
painful to witness. (…) the ordinary yell of rage
which he set up at first, finally changed to a cry(…) uttered somewhat under the breath, andwith a plaintive sound like a moan“Arthur E. Brown (1879)7
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Animal Grief: Perspectives from Anthropology and Activism
Barbara J. King1,
1

William & Mary, U.S.A.

Within evolutionary thanatology, the field of animal-grief studies has grown immensely in the last 15 years.
It sits at a fascinating intersection. Among the media and the animal-loving public there emerges at times areadiness to embrace a romanticized view of other-than-human grief. At the opposite pole, some scientists
cling to hard-core skepticism, wielding the belief that any claims for the expression of animal grief amount tounwarranted anthropomorphism. Meanwhile, scholars of animal grief grapple with how to define andrecognize grief in ways that do not measure expressions of animal emotion only against a human standard.
This work has illuminated the scope and contours of what I believe to be a genuine and profoundly felt grief
in individuals from a variety of mammalian, avian, and perhaps other, species. Based on my own
understandings of animal grief through an anthropological lens, I explore this situation. Then I ask: Can, andshould, this increasing knowledge play a role in animal activism? To what situations in the wild and in
captivity might such knowledge be effectively applied? Would doing so only reinscribe an unwanted speciesist
hierarchy? Or might knowing who in the animal kingdom grieves and how their grief unfolds help us bebetter and do better for animals, who too often pay the price of human exceptionalism?

Are non-human animals likely to have a concept of death?
1Susan Carey1,2Harvard University, 2New York University

My first goal is to explicate what the question of whether animals have a concept of death comes to. Its answer
depends upon what we mean by “having a concept” and what target concept of death we have in mind. I will
characterize the human religious concept of death and the human biological concept of death and will argue
that it is unlikely that any animal has either of these. Animals and prelinguistic infants undoubtedly havemental representations with abstract conceptual content, called systems of core cognition or core knowledge
in the developmental cognitive science literature. Examples include representations of number (systems ofrepresentations with numeral content that are precursors to mathematical knowledge), representations ofintentional and causal agency (a system of representation with agent at its core with content that overlaps andis the precursor to intuitive psychology), and representations of physical knowledge (a system ofrepresentations with the concept object at its core that represents physical interactions among objects). Thesesystems of core cognitive are evolutionarily ancient and are attested early in human infancy in every culture
studied. In support of my argument, I will illustrate the evidence for core cognition of agency, and the lack ofevidence for a biological concept alive in infancy. And I will sketch the evidence that the construction of an
intuitive vitalist biology, the first conceptual system that includes a biological concept of death in childhood,requires conceptual change. I will endorse the suggestion made in some of the abstracts for the present
symposium that the animal concept of death attested in comparative evolutionary thanatology is cessation of
agency. I will pose the question of what precursors to a biological concept of death the representation of
cessation of agency might be shown to include.

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Suicide is Panless: Why Chimpanzees Never Kill Themselves
1

James Anderson1

Graduate School of Letters, Kyoto University, Japan

Chimpanzees in the wild are known to die from various causes, including disease, predation, intra-specificaggression, hunting by humans, and accidents. The literature contains many reports of such deaths. But
despite the many thousands of hours during which chimpanzees have been observed both in the wild and in
captivity, no cases of suicide have ever been witnessed, nor any convincing evidence for deliberate self-killingfound. In fact, the same applies to all nonhuman primates and, I would argue to all nonhuman species. By
contrast, suicide is estimated to account for around one million human deaths worldwide each year; it isconsidered the leading cause of violent death. To understand why suicide appears to occur only in humans,
we need to consider comparative and developmental aspects of death awareness, as well as cognitive processes
that arise from self-awareness, such as self-conscious emotions, mental time travel, and the ability to imagine
alternative future outcomes.

Live or Lifeless? How Chimpanzees Perceive “Death-stimuli“
1André Gonçalves1

Center for the Evolutionary Origins of Human Behavior, Kyoto University, Japan

Humans and many non-human animals divide their world into agents and non-agents. Agents usually haveclear morphological/biomechanical features associated with animacy used to categorize them as such.
According to the animate monitoring hypothesis, animate entities attract more attention than inanimates dueto their potential role as predators or prey in ancestral human environments. Studies show that humans
quickly detect both upright lions and impalas in comparison to their reclining counterparts. Detecting
animacy is a problem many animals face, expected to be widespread in Nature. The perception of dead animals
poses an interesting question since they share both animate and inanimate attributes. Visual-defining featuresof death include inertia, injury, and decay. Using a two-competing stimuli task with eye-tracking, wemeasured chimpanzees’ attention toward images depicting both live and dead animals in realistic backgrounds
to explore if they exhibited preferential-looking patterns towards live versus death-related stimuli. In
experiment1, the stimuli consisted of image-pairs of live and dead animals. Experiment2 had image-pairs ofdead and live infant-carrying behavior in non-human primates. Lastly, experiment 3 consisted of image-pairsof sleeping and dead mammals. Our results suggest a similar animacy bias towards upright live animals versus
dead reclining animals in experiment1; the chimpanzees detected live animals quicker and looked at themsignificantly longer. Experiment2 showed this bias carried through infant primates; chimpanzees looked
longer at live primate infants in comparison to dead ones. Finally, in experiment3, there was no suchdifference in latency or looking durations; sleeping and dead animals were perceived similarly in thisexperiment. The results suggest that indeed chimpanzees show an animacy bias towards life-like stimuli;
however, while we remain agnostic as to whether chimpanzees distinguish sleeping from dead animals inexperimental settings, our sleeping-dead stimuli may be too perceptually similar for such detection in thistask. Other experimental methods involving active choice might reveal such a categorization is within theirgrasp.

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Prospective Studies of Primates' Responses to Death1

Alecia Carter1

University College London, U.K.

The field of evolutionary comparative thanatology is growing rapidly. Despite recent interest in thisrelatively new field of research, there have been few systematic, prospective data. In this talk, I will
discuss prospective observational and experimental research from my lab that queries non-humanprimates' responses to the deaths of others from captive to free-ranging and wild settings. I aim toprovide insight into the breadth and ubiquity of primates' responses to death, their motivations forinteracting with the dead, and evidence for whether they grieve when bereaved.

Equid Responses to Bones and Carcasses
1

Leanne Proops1

University of Portsmouth, U.K.

Myths of elephant graveyards, supported by field experiments and observations, suggest that some nonhuman animals may be attracted to the bones and carcasses of conspecifics long after they have died(Goldenberg & Wittemyer, 2020; McComb et al., 2006). However, recent eye tracking studies withchimpanzees indicate that attraction to the skeletons of conspecifics might reflect a generalized and highly
adaptive drive to affiliative with members of the same species rather than a specific desire to re-visit deadgroup members (Gonçalves et al., 2022). In short, the mechanisms underlying such behaviour remain unclear.
In this talk, I will introduce two ongoing projects that explore domestic horses’ responses to skulls and bones
and wild donkeys’ responses to conspecific carcasses and matched control areas. Equids are an ideal taxonomic
group in which to study this form of mortuary behaviour given that they develop lasting social bonds and canreact strongly to the death of social partners (Dickinson & Hoffmann, 2016). In our domestic horse study,
across three experiments we presented subjects with arrays of 1) a horse skull, cattle skull and pig skull; 2) ahorse skull, horse leg bone and a wooden log; and 3) a horse skull and plastic 3D model of a horse skull, withhalf the skulls presented frontally and half laterally. Our preliminary findings provide insights into thepreferences and discriminative abilities of horses and the role of olfactory versus visual cues in these responses.
Since 2022, in our second study we have positioned video camera traps near ten wild donkey (burro) carcasses
found in the region of Bill Williams River, Arizona, USA. Field notes suggest these carcasses become
important local landmarks where individuals may socialise and establish territorial boundaries. Alongsidethese cameras, we recorded nearby comparable locations, relocated carcass sites, and scent marking posts.
From these data we will be able to calculate visitation rates to determine if carcasses are preferentially visitedand compare detailed activity budgets for donkeys observed at each site to provide insights into the functionof these visits. These are the first systematic studies of responses to bones and carcasses in equids and will help
to elucidate the proximate mechanisms and evolutionary origins of these mortuary behaviours. We also hopethat the establishment of these controlled field studies can provide a blueprint for future studies across species.
Full author list: Leanne Proops1 , Teresa Romero1 , Kate Lewis1 and Erick Lundgren2

Affiliations: 1Centre for Comparative and Evolutionary Psychology, University of Portsmouth, UK; 2Department of Biology, Aarhus University,
Denmark

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Dead Calf Carrying in Asian Elephants
1

Sanjeeta Sharma Pokharel1

Asian and African Area Studies, Kyoto University, Japan

Understanding how animals respond to death is gaining traction among ethologists and conservation
biologists. Thanatalogical responses of highly cognitive and behaviourally flexible species, the Asianelephants, were reported based on opportunistic field observations and publicly available videos*. Similar to
African elephants, Asian elephants showed a range of behaviours, including exploration, visitation, guarding,epimeletic responses and vocalization in reactions to dying and dead conspecifics. Notably, adult female
elephants were observed carrying dead calves, a behaviour rarely observed in elephants. The condition of deadcalves and the mode of carrying raises questions whether elephants truly understand the state of the calves.
The fact that adult females carried dead calves for extended periods, despite the potential hindrance to theirbasic survival activities, such as foraging and movement, hints at associated maternal instincts and suggests
two possibilities either (i) their failure to comprehend the state of ‘death’ and hence, carrying the‘unresponsive’ calves or (ii) the existence of grief-like behaviours in elephants. Further comprehensive studies,including physiological assessments of surviving individuals, are warranted to better understand these
phenomena and to advance the field of comparative thanatology.
*References: Sharma, N., Pokharel, S.S., Kohshima, S. and Sukumar, R., 2020. Behavioural responses of free-ranging Asian elephants (Elephas
maximus) towards dying and dead conspecifics. Primates, 61, pp.129-138. Pokharel, S.S., Sharma, N. and Sukumar, R., 2022. Viewing the rarethrough public lenses: insights into dead calf carrying and other thanatological responses in Asian elephants using YouTube videos. Royal Society open
science, 9(5), p.211740.

Sexual dimorphisms in behavioural responses to a poached conspecific in wild
forest elephants (Loxodonta cyclotis)
Claudia Stephan1Friedrich-Alexander University, Germany

1

The increased attention towards deceased conspecifics in various social animal species seems to be one of themost intriguing conundrums in animal behavior. The factors that might explain the observed behavioral
variation amongst individuals and contextual information that could impact on the latter remain widely
nebulous. Here we analyse forest elephants’ (Loxodonta cyclotis) responses to a poached adult maleconspecific, using remote camera trapping and individual identification during a period of eight months.
After completely avoiding the carcass site for over a week, general interest was high across all age classes.
However, females and males substantially differed in behavioral responses. Males consistently stayed longer
around the remains, showed signs of increased arousal by physically interacting with the bones and thesurrounding vegetation, and bonding behavior. Females, in contrast, were more passively explorative andpreferred to visit the site without their dependent offspring, suggesting that they associated increased risk
with the dead body. Our findings suggest that while females balance potential inclusive fitness risks fromvisiting the carcass against gathering social information, males are mainly driven by the urge to assess
information that might affect their own position in male dominance hierarchies. Forest elephants seem toinfer contextual information associated with death and to adapt their behavioral responses accordingly.
Results reported here highlight the necessity to take intrinsic and extrinsic factors into account wheninvestigating behavioral plasticity towards dead conspecifics. Furthermore, our findings inform adaptive
management strategies with regards to forest elephant conservation efforts.

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The Curious Case of Elephant Burials
1

Nachiketha Sharma1

Institute for Advanced Study, Kyoto University, Japan

Elephants are known to display a wide range of thanatological behaviors from physical interactions, visiting
the carcass and even to carrying the dead. They show deep interest in the remains of dead conspecifics that
may even extend to conspecific skeletons. Of these thanatological interactions, ‘burial’ is one of the mostintriguing and very rarely documented behaviour in elephants. Wild African savanna elephants (Loxodonta
africana) have been, on occasion. observed to partially cover dead conspecifics with branches, leaves or soil.
Some anecdotal reports suggest this species of elephants have buried dead/sleeping humans or even otheranimals. As with dead calf carrying (a behavior observed in two elephant species), a recent study claimed thatwild Asian elephants (Elephas maximus) also engage in burial behaviors similarly to their Africancounterparts; they seem to drag the carcass of calves for burying ‘upside down’ position in ditches. In thistalk, I will present a brief overview of how and why non-human animals bury their dead and through my own
observations, I will discuss the circumstantial evidence to explore the intentionality behind the burial
behaviours of elephants.

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The Thanatosis of Beetles
Takahisa Miyatake1
1

Okayama University, Japan

Thanatosis (or death feigning) is widely observed in animals including mammals, birds, reptiles, fishes, andinvertebrates. Ever since Charles Darwin and Jean-Henri Fabre have shown interest in thanatosis, its adaptive
significance has not yet been proven with sufficient data. Therefore, I examined the conditions under which
thanatosis occurs using the weevil (Cylas formicarius), at first. The weevil feigns death in a state of extreme
rigidity with a characteristic posture that can be discriminated from dead specimens, indicating the weevil
shows consciously feigning dead. Second, a trade-off between flight ability and duration of thanatosis has
been examined using the seed beetle (Callosobruchus chinensis) in lab and field. Third, strains artificiallyselected for shorter (S-strains) and longer (L-strains) duration of thanatosis to measure its inheritance andcorrelated responses using the red flour beetle (Tribolium castaneum). In the field, S-strain type is the
majority and L-strain type is the minority in this beetle species. When cohabiting with a predatory jumping
spider, L-strain beetle was significantly less eaten than S-strain. However, L-strains were at a disadvantage inbreeding because of their lack of movement. This is because they have not met their sexual partners. Beetles
derived from S-strains showed higher expression of dopamine in their brain and higher activity than thesederived from L-strain beetles. When the L-strain beetles were placed on a treadmill to measure locomotor
ability, abnormalities in gait were also found. The duration of thanatosis was shortened by dopamine
supplementation. Transcriptome analysis showed that L-strain beetles had higher expression of dopaminerelated genes in the tyrosine metabolism system and greater variation in human Parkinson’s disease-related
genes compared to S-strains. Although I am aware that my talk is somewhat different from what you may
think of thanatology, insects are clearly consciously performing death feigning. It would be interesting, in myopinion, to know if predators can distinguish between a dead prey and an individual feigning death as a future
work.

From cannibalism to burial: corpse management in termite societies
1

Qian “Karen” Sun1Louisiana State University, U.S.A.

Social insects, such as termites, ants, and bees, frequently face the death of colony members as a consequenceof living in large groups. While dead conspecific individuals are commonly avoided by non-social insects,
social insects employ various corpse management strategies, including removal, burial, and cannibalism. The
behavioral response depends on the nature of the corpse as well as the feeding and nesting ecology of thespecies. The eastern subterranean termite, Reticulitermes flavipes, is a wood-feeding species living in the soil
with thousands to millions of individuals in a colony. In this species, workers cannibalize freshly dead colony
members and bury them when corpses are highly decomposed. This behavioral alteration is modulated bypostmortem changes in the chemical profile of the corpse. Immediately after death, the corpse releases a blendof volatiles, 3-octanone and 3-octanol, which attract workers to retrieve and consume it. The early death cuesenable the colony to recycle nutrients before decomposition occurs. With longer postmortem time, early
death cues diminish, and the accumulation of late death cues (fatty acids, phenol, and indole) indicates corpse
decomposition and elicits burial behavior. Among the fatty acids, oleic acid is a shared death cue among
diverse arthropods, which typically triggers avoidance in non-social species but corpse management in social
species. In R. flavipes, the dynamic change from novel early death cues to conserved late death cues balances
nutritional benefits and pathogenic risks. This postmortem chemical communication highlights uniqueadaptations in termites to their ecological and social niche.

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Death as an indicator of risk in the Scrub Jay
1Teresa Iglesias1

Okinawa Science Institute, Japan

Abstract: Navigating the risk of injury or death is a fact of life for all organisms and exposure to danger canbe managed by assessing and attending to cues of elevated risk. One of the most salient cues of risk is a deadconspecific. I will discuss how the sight of a dead conspecific induces alarm calling and risk-reducing behavior
in a corvid, the western scrub-jay, Aphelocoma californica. Discovery of a dead conspecific elicits vocalizations
that are effective at attracting conspecifics, which then also vocalize, thereby resulting in a cacophonous
aggregation. However, is this reaction simply a surprised response to novel objects that suddenly appear
nearby? How does it compare to seeing a predator? Do they have to recognize the dead individual? Are they
responding to the smell of the conspecific carcass? What about dead heterospecifics? I will describe a series ofexperiments that answer these questions. By exploring the cues that are necessary and sufficient to elicit thesecacophonous aggregations and the subsequent behavioral modification that reduces their exposure to theseareas, we can begin to understand one component of the western scrub-jays’ response to their dead.

The understanding of irreversible non-functionality in Goffin’s cockatoos
Antonio J. Osuna- Mascaró1
1

University of Vienna, Austria

While recognizing when something has permanently ceased to function is of profound importance for humantechnology and social behavior, it remains unexplored in non-human animals. This study investigates theability of Goffin's cockatoos (Cacatua goffiniana) to develop an understanding of irreversible nonfunctionality by using a touchscreen interface. These parrots were able to learn to categorize interactive
elements as irreversibly non-functional after associating them with a particular event, learning to generalize arule by acquiring the necessary understanding of it. The Goffin's cockatoo has shown remarkable tool use andtool making skills in the past, making it an ideal model to study the perception of functionality in non-humananimals. Nevertheless, the understanding of irreversible non-functionality has broader implications, as it isalso a proposed prerequisite for the development of a minimal concept of death.
Full author list: Antonio J. Osuna-Mascaró1, Eleonora Rovegno1,2, Remco Folkertsma1, Susana Monsó2,4, Alice M.I. Auersperg1

Affiliations: 1Comparative Cognition, Messerli Research Institute, University of Veterinary Medicine Vienna, Medical University Vienna,
University of Vienna, Veterinaerplatz 1, 1210 Vienna, Austria; 2Department of Life Sciences and Biotechnology, University of Ferrara, via Borsari
46, 44121 Ferrara, Italy; 3 Department of Logic, History and Philosophy of Science, UNED (Universidad Nacional de Educación a Distancia), Paseo
Senda del Rey 7, 28040 Madrid, Spain; 4 Unit of Ethics and Human-Animal Studies, Messerli Research Institute, University of Veterinary
Medicine Vienna, Medical University Vienna, University of Vienna, Veterinaerplatz 1, 1210 Vienna, Austria

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Abstracts:
Lightning Talks“Histories of Ethiopia, unpolluted by Greek
fripperies, record that if an elephant sees anotherelephant lying dead, it will scoop up some dirtwith its trunk and sprinkle it on the corpse, as if
it were conducting a sacred ritual of some sort,
failing to do so would be sacrilege, once it hasdone its duty, the elephant is free to move on.“Claudius Aelianus (175-255 CE)

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A Concept of Death in Genus Pan: Implications for Human Evolution
1

Katherine H. McLean1

University of Auckland, New ZealandAn understanding of what death and dying entail is termed a concept of death (CoD), and the human CoD
is often viewed as one of the characteristics that distinguishes our species. In this research, I identified ananalogous understanding of death and dying in our closest living relatives—genus Pan. Linguistic frameworks
designed for studying the CoD in human children look for evidence of understanding of several facets ofdeath. I adapted these frameworks for the non-verbal Pan species, systematically analysing written and video
recordings of chimpanzee and bonobo behaviours surrounding death within these new behavioural
frameworks. I identified compelling evidence for the comprehension of several aspects of death, and thus forthe presence of a human-like CoD in chimpanzees and bonobos. This has implications for our own
evolutionary story and raises questions about what makes humans ‘human’.

A case of maternal cannibalism in Macaca fuscata1

Yu Kaigaishi1

Institute for Advanced Study, Kyoto University, Japan

How primate mothers interact with their dead infants has attracted much attention from primatologists. In
particular, the care given to infant corpses has been well documented. In contrast, instances of maternalcannibalism, another striking response to dead infants, are rarely observed and the factors contributing to thisphenomenon remain unclear among primates. Here, I report a new case of stillbirth and subsequent maternalcannibalism which occurred in a free-ranging group of Japanese macaques (Macaca fuscata). In the observedcase, a multiparous female gave a stillbirth in winter, 2022. The newborn appeared premature due to the lackof hair and unformed limbs, and the unseasonal birth. Initially, the mother cared for the newborn by licking
and holding the body, but she suddenly began consuming the entire corpse, except for the parts around thehead. This is the first case of maternal cannibalism by a multiparous mother of free-ranging Japanese macaque.
We compared this case with those previously reported in Old World monkeys, and discussed the possible link
between stillbirth, maternal care and cannibalization towards deceased infants. In order to understand how
primates comprehend the concept of “death” and how their maternal relationships have evolved, it would be
important to focus not only on maternal care toward dead infants but also cannibalism that could occur atthe end of the caregiving.

Response of Irrawaddy dolphins to a dead conspecific1

Amy Jones1.2

Environmental Research Institute, UK. 2Marine Conservation, Cambodia

Cetacean behavior has long attracted scientific attention as humans endeavor to discover what makes thesemammals so emotive and engaging. To date, much of this research has focussed on abundant and widelydistributed cetacean species such as bottlenose dolphins (Tursiops truncatus) and humpback whales(Megaptera novaeangliae). As an endangered and often evasive species, research regarding Irrawaddy dolphin(Orcaella brevirostris) behavior is limited. This study uses data collected by The Cambodian Marine Mammal
Conservation Project, to investigate the behavioral responses of Irrawaddy dolphins towards a deadconspecific. During a routine boat survey of Cambodia's Kep Archipelago, the carcass of an adult female
Irrawaddy dolphin was recovered and attached to the stern of the research vessel and promptly towed to theresearch island for further examination. During this survey, there was a four-fold increase in the number of

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Irrawaddy dolphin groups observed compared to the seasonal average (post-monsoon), in addition to an
atypically positive response towards the research vessel and an atypical increase in the number of behavioral
events observed. These behavioral variations were believed to be in response to the towed dead conspecific.
The authors propose future dedicated research to assess the complexity of wild Irrawaddy dolphin behavior,cognition, and awareness to robustly exemplify the species' apparent sentience and intelligence.

Reaction to Dead Conspecific by European Magpies
1

Olha Brezghunova1

H.S. Skovoroda Kharkiv National Pedagogical University, Ukraine

European magpies (Pica pica) have been observed to respond to dead conspecific by forming an aggregation.
During the study of the behavior (2000-2020) and ecology (2002-2009) of the species in Kharkiv region,
Ukraine, it was only once when such behavior was registered (March of 2003, Siverskyi Donets floodplain).
The "funeral gathering" was formed near an adult magpie that was just-killed by a Goshawk (Accipitergentilis). The Goshawk was no longer near the dead individual and neither the gathering. The birds wereclamoring (sounds of such intensity, a cacophony, were not observed in typical gatherings formed when apredator appears), which attracted more and more individuals (from 10 to 15 birds were present
simultaneously). The birds approached the dead magpie at a distance of up to 30 cm (without touching it). I
discuss different social interactions and responds to dead conspecific among corvids. Some reaction to deadconspecific is known for Pica sp., Corvus sp., Pyrrhocorax sp. Presumably it is a signal to other individuals of
the potential danger.

Do People Believe that Nonhuman Animals Grieve?
1

Heather King1

Rice University, U.S.A.

Do animals grieve? What evidence would convince you they do? Animals’ ‘non-human’ experience of deathand their (in)ability to grieve are frequently represented in the long list of powers which humans have denied
to animals. Despite this, the nature of animal grief remains elusive, with people increasingly willing toattribute emotion, sociality, and indeed, grief, to at least some animals. Regardless of the lack of consensus,
our beliefs about whether animals can comprehend death is a central facet of how we think about animal
minds—and critically, how we conceive of the differences between humans and animals. We take this as ourcritical starting point, and ask to what extent the capacity of animals to grieve is supported by survey
responses. In this lightning talk, I will present the qualitative data gathered from our 2022 survey ‘Do Animals
Grieve?’, led by PI Dr Leanne Proops, in which we collected data from 275 UK-based participants. Our survey
queried participants on various topics related to animal cognition, with a specific focus on various animals’
understanding and experience of death. One common theme in the responses was the discussion of animal
emotion as evidence to suggest animal grief. Such emotionality in animals was then invoked to justify bothwhy, and why not, animals ought to be able to explore the body of close companions after death, withjustifications on either side involving a wish to protect, or refrain from harming, the animal's emotional wellbeing. My talk will summarize the qualitative portion of our survey results, identifying common trends,
discussing anecdotal reports, and analyzing respondent’s storytelling practices.

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Exploring Japanese children's knowledge of death1

Aoi Miyagawa1

Kyoto University, Japan

Human adults understand death to be the irreversible cessation of bodily functions and mental activity. At
what age is this understanding formed? Researchers have reported that understanding of death develops by
the age of 7 years and reaches the same level of understanding as adults around the age of 10-12 years (Speeceand Brent, 1984, 2007; Slaughter, 2005). However, it has been suggested that understanding of human deathmay be influenced by a person's cultural background (Brent et al, 1996). The present study addressed possible
cultural influences by comparing Japanese children’s understanding of death with that of Western children.
It also examined the influence of personal experiences on understanding of death, by asking parents about
their children's previous exposure to death.

Responses to the corpse observed in wild stump-tailed macaques in Thailand
Aru Toyoda1,2,3,4

1

Japan Society for the Promotion of Science, Japan Monkey Centre, National Primate Research Center of Thailand, Wildlife Research Center of Kyoto University
2

Necrophilic behavior (attempted copulation with corpses) is rarely documented in nonhuman primates,
especially in wild populations. This presentation shows the first recorded case of necrophilic behavior in wildstump-tailed macaques in Thailand. Six groups totaling over 460 individuals, were identified and habituatedfor observation. The corpse of an adult female, identified as TNG- F11, was discovered and observed directly
and via camera trap for three days. Although the cause of death remained undetermined, no significant
physical injuries were observed. During the three-day observation period, three different males attempted tocopulate with the corpse. Notably, these males included not only those from the deceased female&#39;s groupbut also from different groups. These observations suggest that certain cues emitted by the corpse, along with
its nonresistant/passive state, may have triggered these behaviors in the males. Given the rarity of necrophilic
behavior reports in non-human primates, our findings offer new insights into such behaviors and contributeto the broader understanding of comparative thanatology. In this presentation, we report not only the detailsof these necrophilic behaviors, but also newly observed cases and previously observed cases of reactions tocorpses, including previous cases of carrying dead infants.
Full author list: Aru Toyoda 1,2,3,4 , André Gonçalves 5 , Tamaki Maruhashi 6 , Suchinda Malaivijitnond 3,7 , and Ikki Matsuda 4,8,9,10
1 Japan Society for the Promotion of Science 2 The Japan Monkey Centre, Inuyama, Aichi 484-0081 Japan 3 National Primate Research Center of Thailand 4 Wildlife Research Center of Kyoto University 5 Section of Cognitive
Neuroscience, Center for the Evolutionary Origins of Human Behavior, Kyoto University 6 Musashi University 7 Department of Biology, Faculty of Science, Chulalongkorn University 8 Chubu Institute for Advanced Studies, Chubu
University 9 Chubu University Academy of Emerging Sciences 10 Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah

Chimpanzee behavioural reactions to putrescine, a chemical cue of death1

Hanling Yeow1.2

Max Planck Institute for Evolutionary Anthropology, Germany,2Kyoto University, Japan

Studying animal reactions to dead and dying conspecifics can inform us how much they understand and areaware of death. In chimpanzees, dead infant carrying is a puzzling behaviour because decaying corpses may
carry the risk of disease transmission. In this study, putrescine, an unpleasant-smelling chemical produced
when bodies decay, was presented to chimpanzees along with the dead body of a bird, and their reactions wererecorded. Chimpanzees avoided putrescine but not the dead bird, suggesting that they do not show aversion
towards corpses by themselves, but may show aversion towards chemical cues of decay. Dead infant carrying
may be a naturally occurring behaviour in the early stages after death, when cues of disease are not yet present.
However, cases of prolonged dead infant carrying behaviour occur despite the aversion to cues of potential
infection from the decaying corpses, and further research will be required to understand why this happens.

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Schedule“Instead of being scared away by the report of
the gun, leaving their wounded or deadcompanion to his fate, they show the greatest
anxiety or sympathy for him, uttering cries of
distress, and plainly proving that they wish torender him assistance by hovering over him…“John Selby Watson (1870)

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20th Thursday

21st Friday

The thumb and the trunk:
Experimental and observational insights into non-humanprimates and elephants' death awarenesss

Silent antenna, rustling feather:
Lessons from the lab to the field, from insect thanatosis
to avian thanatology

09:00 - 09:15 – INTRO & REGISTRATION

09:00 - 09:15 – INTRO & REGISTRATION

Keynote Talk:
09:20 - The Child's Construction of a Concept of Death
Susan Carey, Harvard University*

Keynote Talk:
09:20 - Animal Grief:
Perspectives from Anthropology and Activism
Barbara J. King, William & Mary*

10:05 - 12:05 - SESSION 1: Perceptive Primates
Chair: André Gonçalves
10:05 - Suicide is Panless:
Why Chimpanzees Never Kill Themselves
James Anderson, Kyoto University
10:45 - Prospective Studies of Primates' Responses to Death
Alecia Carter, University College London
11:25 - How Chimpanzees Perceive Death-stimuli
André Gonçalves, Kyoto University
12:05 - 13:30: Lunch Break
13:30 - 14:25 - SESSION 2: Lightning Talks I*
Chair: Sarv Dashti

10:05 - 11:25 - SESSION 4: Imperative Invertebrates
Chair: Sanjeeta Pokharel
10:05 - The Thanatosis of Beetles
Takahisa Miyatake, Okayama University
10:45 - From Cannibalism to Burial:
Corpse Management in Termite Societies
Qian Sun, Louisiana St. University*
11:25 - 12:25 - SESSION 5: Lightning Talks II*
Chair: Nahoko Tokuyama
12:25 - 13:30: Lunch Break

14:30 - 16:50 - SESSION 3: Elusive Elephants-Equids
Chair: Alecia Carter
14:30 - Equid Responses to Bones and Carcasses
Leanne Proops, University of Portsmouth
15:10 - Dead Calf Carrying in Asian Elephants
Sanjeeta S. Pokharel, Kyoto University
15:50 - Sexual Dimorphisms in Behavioural Responses to a
Poached Conspecific in Wild Forest Elephants
Claudia Stephan, Friedrich-Alexander University*
16:20 - The Curious Case of Elephant Burials
Nachiketha Sharma, Kyoto University

13:30 - 14:30 - SESSION 6: Brilliant Birds
Chair: Nachiketha Sharma
13:30 - Death as an Indicator of Risk in the Scrub Jay
Teresa Iglesias, Okinawa Inst. Sci. Tech.
14:10 - The Understanding of Irreversible Nonfunctionality in Goffin's Cockatoos
Antonio Osuna-Mascaró, Vetmeduni*
14:50 - 16:20 - WORKSHOP
14:50 - What Could Future Thanatology Look Like?
Alecia Carter, University College London*Online Talks

2020th Thursday
Lightning Talks I
13:30-14:18

13:30 - A case of maternal cannibalism in Macaca fuscata
Yu Kagaishi, Kyoto University
13:42 - Reponses to the corpse observed in Wild Stumptailed macaques in Thailand
Aru Toyoda, Japan Monkey Centre
13:54 - Reaction to dead conspecific by European Magpies
Olha Brezghunova, H.S. Skovoroda Kharkiv National
Pedagogical University*
14:06 - Response of Irrawaddy dolphins to a dead conspecific
Amy Jones, Environmental Research Institute*

21th Friday

Lightning Talks II
11:25-12:23
11:25 - Do People Believe that Nonhuman Animals Grieve?
Heather King, Rice University*
11:47 - A Concept of Death in Genus Pan: Implications for
Human Evolution
Katherine H McLean, University of Aukland*
11:59 - Exploring Japanese children's knowledge of death
Aoi Miyagawa, Kyoto University
12:11 - Chimpanzee behavioural reactions to putrescine, achemical cue of death
Hanling Yeow, Max Planck Institute for
Evolutionary Anthropology*Online Talks

212AffiliatedInstitutions“Sometimes a herd of large and small dolphinshas been seen together, and two of these having
left, appeared soon after, supporting andcarrying on their back a small dead dolphin, thatwas ready to sink, as if in pity for it, that it
might not be devoured by any other wild
creature“
Aristotle (384-322 BCE)

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In collaboration with:

And partnership of:Funding:

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EvoThan2024 was organized by:André Gonçalves - Nachiketha Sharma
Sanjeeta Pokharel - Sarv Dashti
Alecia Carter - Nahoko Tokuyama

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