Picture is made by “AndreaMosaic”©Hiroya TAKIYAMA

Date

September 22nd (Sat)-24th (Mon), 2018
※Monday is a national holiday

Venue

Science Seminar House,
North Campus, Kyoto University

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th

The 10 International Symposium on Primatology and Wildlife Scienceth- The 10 Anniversary of Wildlife Research Center PROGRAM
Day 1 September 22nd (Sat)Time(min) Title

Speaker

13:00-13:10

13:10
|
14:00

(10) Opening Remarks

Tetsuro Matsuzawa PWS Program Coordinator

WRC 10th Anniversary Special Event I

Chair: Satoshi Hiratai

O-01 (10) Introduction to the WRC 10th Anniversary Special Event Satoshi Hirata

Wildlife Research Center, Kyoto
University

O-02 (10)Attentional and physiological response toward others’
injury and pain among chimpanzees

Yutaro Sato (L2)Kristin Havercamp Wildlife Research Center, Kyoto(L3)
University

O-04 (10) Lateral position preference in feral horses

Sota Inoue (L3)

Utilizing The Great Ape Information Network (GAIN) to understandO-05 (20)Modeling Habitat Suitability for Yunnan Snub-nosed
Monkeys (Rhinopithecus bieti) in Laojun Mountain

Jie Liu (L5)

Wildlife Research Center, Kyoto
University

O-06 (10)Assessment of the current impact of mountain gorilla
ecotourism in Bwindi National Park, Uganda

Raquel Costa (L4)Primate Research Institute,
Kyoto University

O-07 (10)Male-male relationships of chimpanzees in Kalinzu forest:
Aggressive behavior and party composition

Shohei Shibata(L2)

Primate Research Institute,
Kyoto University

Jie Gao (L3)Primate Research Institute,
Kyoto University

Andre Goncalves

Primate Research Institute,
Kyoto University

O-09 (10)(15)

Death Among Primates: A critical review of non-humanprimate interactions towards their dead and dying<Coffee Break>Invited talks by new PWS members

Chair: Yuko Hattori

O-10 (15) Read my lips: when the red gets redder
Images speak louder than numbers? Evaluation of liver function

O-11 (15) by ultrasound and serum biochemistry in owl monkeys
15:15
|
16:45

O-12 (15)Sensitivity to Bitter Molecule Phenylthiocarbamide(PTC) in Four Species of Sulawesi Macaques

Lucie Rigaill

Primate Research Institute,
Kyoto University

Sayuri Takeshita

Primate Research Institute,
Kyoto University

Kanthi Widayati

Department of Biology, Bogor
Agricultural University, Indonesia

O-13 (15) How to rejuvenate the super-aged society

Kaoru Sekiyama

O-14 (15) Working for International Organizations

Eriko Kawai

O-15 (15)Working for an international organization:communication skill

Yoshihiro Kawai

Enrichment studies and an invitation to the 14th ICEE

16:45
|
17:30

17:30-18:30

Wildlife Research Center, Kyoto
University
Chair: Lira Yu

O-08 (10) The understanding of body structures in chimpanzees

15:00-15:15

Wildlife Research Center, Kyoto
University

O-03 (10) longevity and mortality patterns of chimpanzees in Japan

Field and laboratory studies on primate species I

14:00
|
15:00

Affiliation

Registration

12:00-13:00

Graduate School of Advanced Integrated
Studies in Human Survivability, Kyoto
University
Graduate School of Advanced Integrated
Studies in Human Survivability, Kyoto
University

Graduate School of Management,
Kyoto University

Chair: Yumi Yamanashi

O-16 (15)Filling the gap: an update from the Kyoto City Zoo andan invitation to the ICEE2019

O-17 (10)Verification of the effectiveness of environmental
Momoko Oka (L2)enrichment and the influence by visitors in captive tigers

O-18 (10)Primate Research Institute,
Salivary alpha-amylase enzyme as a non-invasive biomarker
Nelson Broche (L2)of acute stress in Japanese macaques (Macaca fuscata)
Kyoto University

O-19 (10)“Bar hanging” behavior: a look into a potentiallyculturally propagated behavior and its relation to stress
Poster session/ PWS exam

2

Yumi Yamanashi

Josue Pastrana(L4)

Kyoto City Zoo
Wildlife Research Center, Kyoto
University

Primate Research Institute,
Kyoto University

Day 2September 23rd (Sun)Time(min) Title

Speaker

WRC 10th Anniversary Special Event II

9:00
|
9:50

9:50-10:00

Chair: Shiro Koshima

O-20 (10)Studies on ecology and behavior for wildlife
conservation

Shiro Koshima

Wildlife Research Center, Kyoto
University

O-21 (20)

Eavesdropping on elephants: How and what do Asianelephants talk with each other?

Nachiketha
Sharma (L5)

Wildlife Research Center, Kyoto
University

O-22 (20)

Short-term gut microorganism dynamics induced bygumnivorous feeding for lesser slow loris

Kei Matsushima(L5)

Wildlife Research Center, Kyoto
University(10)<Group Photo>

Penguins, in full color

Chair: Andrew MacIntosh

O-23 (30) Marching with Penguins

10:00
|
11:30

Affiliation

O-24 (20)Just like clockwork? On the significance of periodic
penguins

O-25 (20) Jelly in the Belly: Why do penguins eat jellyfish?

O-26 (20)Solving the mystery of female-biased stranding in
Magellanic penguins

Lloyd Davis

University of Otago, New ZealandAndrew MacIntosh

Primate Research Institute,
Kyoto University

Jean Baptiste
Thiebot

BioScience Group, National
Institute of Polar Research,Tokyo
Research and Development Center for

Takashi Yamamoto Data Assimilation, The Institute of
Statistical Mathematics, Tokyo

11:30–13:00(90)<Lunch Break>

13:00
|
16:00

(180)

Workshop on Science Communication

16:00-16:30(30)

Wiebke Finkler and Llyoyd Spencer Davis<Coffee Break>Field and laboratory studies on wild mammals

Chair: Takushi Kishida

Is a Bonobo Older Brother or just Meat for Bongando
Graduate school of Asian andO-27 (20) Ethnic Group? -Focusing on Local Perceptions towards Aya Yokotsuka (L5) African Area Studies, Kyoto
Bonobos in DRCongoUniversity
16:30
|
17:30

17:30-18:30

18:30–

O-28 (20)Social and Sexual Development up to Natal Dispersal in
Kazuya Toda (L5)
Female Bonobos

O-29 (15)

Did life in trees drive the adaptive evolution of taste inkoala and primates?

Takashi Hayakawa Japan Monkey Centre / PRI

Poster session/ PWS exam

Get-together Party

3

Primate Research Institute,
Kyoto University

Day 3September 24th (Mon)Time(min) Title

9:00-9:15(15)

SpeakerPoster award

9:15–9:45

Chair: Lucie Rigaill(10) Presentation 1

Award Winner 1(10) Presentation 2

Award Winner 2(10) Presentation 3

Award Winner 3

Field and laboratory studies on primate species II

Chair: Yuta Shintaku

Chimpanzees’ behaviour in the prosocial choice
O-30 (15) paradigm depends on the position of the token in thefeeder
09:45
|
10:30

O-31 (15)Male Reproductive Success in Bornean Orangutans(Pongo pygmaeus)

Research, Conservation and Management of the
O-32 (15) Proboscis Monkey Population in Klias Peninsula,
Sabah, Malaysia

Macjei Trojan

Department of Ethology andComparative Psychology, Nicolaus
Copernicus University, Poland

Tomoyuki Tajima

Graduate School of Science,
Kyoto University

Henry Bernard

Institute for Tropical Biology andConservation, University
Malaysia Sabah, Malaysia

WRC 10th Anniversary Special Event III

10:30
|
11:50

Chair: Miho Murayama

O-33 (10) The way forward of wildlife research

Miho Murayama

Wildlife Research Center, Kyoto
University

The Arctic Tundra & the African Savannah: Common
O-34 (15) Conservation Conundrums Confronting Caribou andGiraffes

Fred Bercovitch

Save The Giraffes, San Antonio,
TX, USA

O-35 (15) Culture, social learning and death in Japanese monkeys Claire Watson

Wildlife Research Center, Kyoto
University

O-36 (15) Bats: How to study animals you cannot see

Wildlife Research Center, Kyoto
University

O-37 (15)
11:50-12:00

David Hill

Evolution of the loss of olfaction in whales provides new
Takushi Kishidaperspectives on conservation of the marine amniotes(10) Closing remarks

Wildlife Research Center, Kyoto
University

Tetsuro Matsuzawa PWS Program Coordinator

12:30-13:30 Lunch seminar (Report on Ministry of the Environment " Wildlife Conservation")

14:00
|
17:30

Affiliation

PWS Faculty meeting

Conserv' Session
Last of the Longnecks

4

Chair: Seiko Fukushima

Poster Session

P-01

Title

Presenter

Effectiveness of species recognition depends on body size in the sailfin

Elio Borghezan

tetra: a bridge between mate choice and ontogeny
P-02

Seasonal changes in bats occurrence and activity in the Janówek Fort

Judyta Gulatowska(Masovian District, Central Poland)
P-03

Genetic diversity analysis of the Ryukyu flying fox

Yuto Taki

P-04

Vocalizations of a new population in an avian ring species & research

Annegret Moto Naito

plan for the conservation genetics of the Japanese golden eagle
P-05

Do primates yuck? Pathogen and parasite avoidance behaviors inCécile Sarabian

Papionini and Hominini
P-06

Networks and the role of incomplete data in assessing the impacts of

Zhihong Xusocial structure on parasite flow
P-07

Where do female bonobos migrate?: Implication from mitochondrial

Shintaro Ishizuka

DNA analysis
P-08

Ordering strategies of three-choice task by free-ranging Japanese

Shenwen Xu

macaques
P-09

Food transfer occur between closer individuals?

Makiko Take

P-10

Waiting as Infant Care in Wild Chimpanzees

Hiroko Sakuragi

P-11

Social grooming in rescued chimpanzees: how early social disruption

Evelina Daniela

and time of rehabilitation affect sociality?

Rodrigues

Reaction of a group of chimpanzees to the death of the oldest female in

Julia Sikorska

P-12

the City Zoological Garden of Warsaw
P-13

Spatial representation of age in chimpanzees

Yuri Kawaguchi

P-14

Eye Contact During Gestural Communication in Captive Chimpanzees

Hanling Yeow

and Bonobos
P-15

Development of Combinatory Manipulation in Great Apes and Humans:

Misato Hayashi

Implication for Action Patterns in Tool Use
P-16

An ongoing research:

Yena Kim

familiar vs. unfamiliar human face recognition in chimpanzees
P-17

Chimpanzee’s categorical classification of the sound: The voice ofchimpanzee vs pure tone

5

Hiroya Takiyama

P-18

Informing conservation efforts through quantitative assessments ofMaegan Fitzgerald

habitat suitability for chimpanzees in the Greater Nimba Landscape,Guinea, Africa
P-19

Characterization of bitter taste sensitivity of four species of Sulawesi

Xiaochan Yan

Macaques
P-20

Sedative and Antinociceptive Effects of S(+)-Ketamine and Midazolam

Vanessa Grisor Fentanyl and Droperidol in Black-tufted-ear marmosets
P-21

Inter-group relationships of feral horses (Equus caballus) in Serra

Tamao Maeda

D’Arga, Portuga
P-22

Investigating the social behaviour and structure differences between

Pandora Francisca

single and multi-stallion groups of feral horses

Costa Barão Pinto

P-23

Do female feral horses (Equus caballus) choose their herd?

Sakiho Ochi

P-24

A study plan about feeding behavior in captive giraffes

Anna Kawakita

P-25

Using dynamic animal social networks to improve the study of infectious

Kenneth Keukdisease transmission

6

O-1

Introduction to the WRC 10th Anniversary Special Event

Satoshi Hirata1*
1

Wildlife Research Center, Kyoto University, Kyoto, Japan*satoshi.hirata.8z@kyoto-u.ac.jp

In my talk I will start by giving a brief introduction to this series of special events
during the symposium, the WRC 10th Anniversary Special Event, followed by aprogress report of my own activities. What are human beings? What are minds thatquestion about human beings? Starting with such questions, I have entered into theacademic world to study nonhuman primates from comparative cognitive perspectives.
The main target of my research is the chimpanzee, one of the closest living relatives ofhumans, since my graduate course. I would like to explore the evolution of human mind
through the investigation of social intelligence in great apes. As the Director, I havebeen conducting works at Kumamoto Sanctuary, Wildlife Research Center, Kyoto
University, since 2015. Kumamoto Sanctuary houses the largest population of captive
chimpanzees in Japan, and the only group of bonobos in Japan. Our work at Kumamoto
Sanctuary include experimental cognitive studies with the chimpanzees and bonobos,
naturalistic observation of their behavior from ethological perspectives, and monitoring
their health and stress from an animal welfare perspective. Integration of research andanimal welfare is the important direction of our organization. In addition to the studies
with great apes, I have also started a project to study wild horses in Portugal. In ourstudy, use of drones has opened a new window to investigate group behaviors ofmammals. Predator-prey relationship between horses and wild wolves is one ofinteresting research topics. Following my talk, some students of mine will provide
progress reports about their own studies with great apes and horses.

7

O-2

Attentional and physiological response toward others’ injury and pain
among chimpanzees
Yutaro Sato1*, Fumihiro Kano2, Satoshi Hirata1
1
2

Wildlife Research Center, Kyoto University, Kyoto, Japan

Institute for Advanced Study, Kyoto University, Kyoto, Japan*sato.yutaro.76w@st.kyoto-ac.jp

Nonhuman animals are known to express negative emotions when observing others’
negative emotional expressions, a behavior often regarded as an expression of empathy.
It is also known that humans experience negative emotions (e.g. disgust or pain) whenobserving others’ injury even without observing negative emotional expressions. Thisbehavior is likely not only based on automatic reflection of others’ emotional states but
also based on their understandings about the cause of distress. Yet, this has not been
examined in nonhuman primates. Chimpanzees (Pan troglodytes) are observed toinspect groupmates’ injuries and are also known to console those being distressed. In
this study, we tested six chimpanzees living in Kumamoto Sanctuary, Kyoto University(one adult male, three adult females, two juvenile females), to examine how they react
to others’ injury and pain. First, we used eye-tracking to examine if chimpanzees
spontaneously attend to injured conspecifics more strongly than non-injured ones. We
confirmed that they did so; and also that they did not merely react to conspicuous
features of injury (e.g. red color). Then, we used thermal imaging to examine if
chimpanzees physiologically respond to others’ injury. Previous studies have confirmedthat drop in nasal temperature is a signature of arousal, especially that of negative
valence. We conducted two experiments presenting to them either the still images of
injured conspecifics or movie clips of conspecifics being injected. Chimpanzees did not
distinguish images of injured conspecifics (similar to those used in the eye-tracking
experiment) from those of uninjured in their thermal signatures. We assumed that thepresented stimuli were not realistic or arousing enough for the chimpanzees. We thendemonstrated an injury in real-life; a familiar human experimenter used a magic trickshowing a damage on the skin and running blood; the control action was washing handswith soaps and water. Chimpanzees, especially adult females, exhibited larger
temperature drop in response to the fake injury than to the control stimulus. We thenpresented a condition that is potentially more demanding to understand; a familiar
experimenter stubbed a (fake) thumb with needles without running blood; the control
was stubbing a wood piece with needles. Chimpanzees did not distinguish this stimulus
from the control as a group. Our conclusion is that chimpanzees do experience negative
emotions when observing others’ injury without directly observing others’ emotional
expressions, indicating that they understand the cause of others’ distress as well as
experiencing an empathy. However, it was also clear that not all chimpanzees showed
this tendency; in particular, we found that a male and juveniles did not respond asstrongly as adult females did. Such individual differences might reflect modulation of
empathy by social and experiential factors, which needs to be examined in futurestudies.

8

O-3

Utilizing The Great Ape Information Network (GAIN) to understandlongevity and mortality patterns of chimpanzees in Japan

Kristin Havercamp1
1

Wildlife Research Center, Kyoto University, Japan

kristin.havercamp.24z@st.kyoto-u.ac.jp

Nationwide databases containing individual life history details are important for the
successful management and monitoring of captive animals, but they are rarely available
as open-access. The Great Ape Information Network (GAIN) is a unique example ofone such database. Since the arrival of the first chimpanzee estimated to be in 1921,
1,017 chimpanzees have either lived or currently live in Japan and their individual
details have been recorded and subsequently collated by GAIN. Using GAIN data andtwo softwares, PopLink and PMx, I am creating a national chimpanzee studbook andinvestigating their life history patterns, beginning with the calculation of longevity andmortality rates. Goals of this project are to expand our relatively limited knowledge ofcaptive chimpanzee life expectancy, and more broadly to better understand the life
patterns of chimpanzees in Japan and how they compare to findings from other captive
studies and the wild. I will also discuss the progress of my upcoming project to monitor
captive chimpanzee sleep at Kumamoto Sanctuary.

9

O-4

Lateral position preference in feral horses

Sota Inoue1*, Monamie Ringhofer2, Renata Mendonça3, Shinya Yamamoto2, Satoshi Hirata1
1
2

Wildlife Research Center, Kyoto University, Japan

Institute for Advanced Study, Kyoto university, Japan
3

Primate Research Institute, Kyoto university, Japan*sota.inoe@gmail.com

Lateralization is one of the broadest issue regardless the behavioral types and species.
In mother-infant interactions in many mammals, it has been reported that foals show thepreference to hold their mother in the left eye view field. However, there are few studies
regarding the foals’ preference regarding the spatial positioning to nearest neighbor. In
addition, the adults’ preference has not been reported as well. We made hypothesizes; 1)
individuals will show a lateral spatial positioning preference to nearest neighbors, 2)
adults have the lateral preference in the cross-resting behavior. Foals showed thepreference to hold nearest neighbors in their left eye’s view field in the close distance tofoals. In the case of adults, it depended the positioning of nearest neighbor. If nearest
neighbors were located in the rear area, the standard individual preferred to hold them inthe left eye’s view field. If they were in the front, preferred to hold them in the righteye’s view field. Furthermore, in the population level, adults also showed using the left
eyes during cross-resting behavior. The present study showed the laterality regarding
positioning of nearest neighbors in feral horses.

10

O-5

Modeling Habitat Suitability for Yunnan Snub-nosed Monkeys(Rhinopithecus bieti) in Laojun Mountain

Jie LIU1*, Maegan FITZGERALD1, Hahong LIAO2, Satoshi HIRATA1, Tetsuro MATSUZAWA3
1

Wildlife Research Center, Kyoto University, Kyoto, Japan
2

The Nature Conservancy, Lijiang Office, China

3

Institute for Advanced Study, Kyoto University, Kyoto, Japan*liujie871019@gmail.com

Habitat evaluation procedures are used to quantify the value of land as habitat for a
species. In this study, we analyze environmental variables hypothesized to influence
habitat suitability for Yunnan Snub nosed monkeys (Rhinopithecus bieti) and map the
distribution of suitable habitat across the study area and adjacent areas. The study was
carried out in three ways for data collection: 1) 72 trap cameras, 2) drone observationand 3) tracking on foot. Spatial analysis combining the GPS data was conducted toinvestigate home range change of this monkey group. We collected 958 occurrence
points from 2010 and 2017 in Laojun mountain. Predictor variables were generatedusing ArcMap and R software. We prepared 34 environmental variables at 30 m spatial
resolution. Maxent software was used to analyze environmental variables contributing
to suitability. In general, using satellite remote sensing and GIS, we modeled the
distribution of suitable habitat for Yunnan snub-nosed monkeys in the Jinsichang area of
the Laojun Mountains of China. The habitat suitability result of synthetic effect ofmuti-variables. The environmental variables contributing most to habitat suitability,
measured by permutation importance, were mean diurnal range, precipitation during thewettest quarter of a year, average annual precipitation, NDVI, wetness and aspect.
Habitat suitability model provide important methods and result about potential current
species distribution which is essential for the implementation of conservation actions.

11

O-6

Assessment of the current impact of mountain gorilla ecotourism in
Bwindi National Park, Uganda

Raquel Costa1,2*, Misato Hayashi1, Michael A. Huffman1, Gladys Kalema-Zikusoka3, Lilly
Ajarova4, Fred Bercovitch5,6 and Masaki Tomonaga1
1Primate Research Institute, Kyoto University, Japan
2
3

4

Primate Cognition Research Group, Portugal

Conservation Through Public Health, Uganda

Chimpanzee Sanctuary & Wildlife Conservation Thrust, Uganda
5
6

Save the Giraffes, USA

Wildlife Research Centre, Kyoto University, Japan*raquelberingei@gmail.com

Ecotourism is an emergent and controversial topic in the field of wildlife
management. Ecotourism plays a significant role in the conservation of mountaingorillas (Gorilla beringei beringei). Despite the growing number of tourists visiting
mountain gorillas and an increasing number of habituated groups, very little behavioral
data has been collected on the potential impacts of ecotourism on these wild populations.
The present study examines how interactions with human tourists influence gorilla
behaviour. For this study, we collected behavioural data (focal and scan sampling)
before, during and after tourist visits over a 6-month period for (December 2017-August2018) one habituated group (15 individuals) in Bwindi Impenetrable National Park,
Uganda. Kruskall Wallis tests are used to compare pre-contact, contact and post-contact
periods. Preliminary analysis shows that “Human directed behaviour” increases 3.5
times during the tourist visit (Kruskal- Wallis=10.029, P=0.027 Bonferroni correction)and is present only when tourist are within 3 meters of the gorillas. Moreover, groupsocial dynamics are affected by close distance to humans - affiliation, agonism andsubmission behaviours increase within 3 meters from the tourists. Further analysis isnecessary, but the present results caution a disturbance in the gorilla`s behaviour related
to the visit tourist. The problems are most pronounced when people get too close to thegorillas, in violation of standard rules.

12

O-7

Male-male relationships of chimpanzees in Kalinzu forest: Aggressive
behavior and party composition

Shohei Shibata1*, Chie Hashimoto1, Takeshi Furuichi1
1

Primate Research Institute, Kyoto University, Aichi, Japan*

Shibata.Shouhei.73c@st.kyoto-u.ac.jp

Chimpanzees and bonobos are known for large differences in male aggression inspite of their highly close phylogenetic relation to each other. While lethal aggressionsof male chimpanzees are observed in both intra-group and inter-group relationships,
such kind of behavior is not observed in bonobo society. Investigating their aggression
could give us important hints to understand human aggression.
Both species have differences in degree of fission-fusion tendencies. While
chimpanzees form parties which vary in size and composition, bonobos form a cohesiveand stable large party that includes most of the community members. How important is
the fission-fusion system for their society and what kind of factors shape the differencein cohesiveness of the party? One possibility is that this difference reflects differences
in male aggression. Large parties include many individuals could create social tension
conditions that evoke aggressive interaction among individuals, which may be a leading
factor in chimpanzees dividing into small parties in order to avoid unnecessary
aggressive interactions. In this study, I focus on the relationships between party size,
aggressive behavior, and hormonal secretion of male chimpanzees.
From September to October 2017, from February to April and from June to
September 2018, I observed male chimpanzees and collected urine and fecal samples in
the M group, at Kalinzu Forest Reserve. I conducted focal animal sampling andrecorded social and aggressive behaviors of males as well as the party composition inwhich the focal animal was present. We found that the number of males in a party arelarger in the presence of females showing maximum sexual swelling. In both conditions
where females showed the absence and presence of maximum sexual swelling, thefrequency of male aggressive behaviors increased along with increments of males to theparty. In a party which includes a few number of males, aggressive behaviors wererarely observed. These results suggest that male chimpanzees are dispersing to avoid
expressing and receiving intense aggression. In the next step of our study, we willconduct hormonal analysis of cortisol and testosterone in order to investigate the focal
subject’s physiological markers in both conditions of the absence and presence offemale showing maximum sexual swelling.
Keywords: chimpanzees, party composition, aggression, stress

13

O-8

The understanding of body structures in chimpanzees

Jie Gao1*, Masaki Tomonaga1
1Primate Research Institute, Kyoto University, Inuyama, Aichi, Japan*

gao.jie.87c@kyoto-u.jp

The recognition of bodies of conspecifics’ and other species’ is vital for animals tolive. In this study, we investigated the understanding of body structures in chimpanzees,
both of chimpanzee bodies, and of bodies of other species. Previous studies have foundthat chimpanzees show the body inversion effect to intact chimpanzee bodies, but not toother objects, including houses. It suggests that they use configural processing for theirown bodies. We used scrambled chimpanzee bodies and other species’ bodies withdifferent postures to test whether they use the same way to perceive them as with intact
chimpanzee bodies by examining the inversion effect. For their own species, they
showed no inversion effect for bodies with abnormal body part arrangement, andshowed the inversion effect for bodies with abnormal body part proportion, but theeffect was smaller than for intact bodies. It suggests that chimpanzees have theknowledge of their body part arrangement and body proportions. For other species, they
showed the inversion effect to bipedal humans (familiar but different postures from
chimpanzees) and monkeys (familiar and similar postures as chimpanzees), and to
crawling humans (unfamiliar but similar postures) and horses (unfamiliar but similar
postures). It suggests that chimpanzees’ configural body processing may rely on bothvisual and embodied expertise.

14

O-9

Death Among Primates: A critical review of non-human primate
interactions towards their dead and dying

André Gonçalves1
1

Primate Research Institute, Kyoto University, Japana.gnclves@gmail.com

For the past two centuries, non-human primates have been reported to inspect, protect,retrieve, carry or drag the dead bodies of their conspecifics and, for nearly the sameamount of time, sparse scientific attention has been paid to such behaviours. Given that
there exists a considerable gap in the fossil and archaeological record concerning how
early hominins might have interacted with their dead, extant primates may provide
valuable insight into how and in which contexts thanatological behaviours would haveoccurred. This review expands and builds upon previous attempts to evaluate the bodyof knowledge on the subject. First, we outline a comprehensive history of comparative
thanatology in non- human primates, from the earliest pre-ethological accounts to thepresent, bringing to light the interpretations of previous researchers and theircontributions to the field of primate thanatology. We also show that many of the typicalbehavioural patterns towards the dead seen in the past are consistent with the ones
observed today. Second, we review the recent evidence of thanatological responses andorganise it into distinct terminologies: direct interactions (physical contact with thecorpse) and secondary interactions (guarding the corpse, vigils and visitations). We also
provide a critical evaluation of the behavioural and emotional aspects of these responses
in infants, adults and non- conspecifics, pointing towards its form and function. Third,
we provide an integrative perspective of primate thanatology, bringing together different
fields of research and their recent advances to clarify how thanatological behaviours
would have emerged on the evolutionary, sensory/cognitive, developmental andarchaeological spectrum. For that purpose, we not only suggest an evolutionary timeline
for thanatological responses typical to primates, but also propose an integrated model ofLife-Death Awareness, whereupon neural circuitry dedicated towards detecting life
(animate agency, intentional agency, mentalistic agency) works with a matching systemthat interacts with it on a decision-making level (animate/inanimate distinction,living/dead discrimination, death awareness). Finally, we discuss the importance ofdetailed reports on primate thanatology and propose different empirical avenues to shedfurther light on this topic.

15

O-10

Read my lips: when the red gets redder

Lucie Rigaill1*
1

Primate Research Institute, Kyoto University, Inuyama, Aichi, Japan*rigaill.lucie.2c@kyoto-u.ac.jp

Across human cultures, red is often associated with love and fertility. Men, fromdifferent populations, found women wearing or surrounded by red more attractive, andwomen, subconsciously, are also more likely to wear red ornaments when they are mostfertile, i.e., around ovulation. Several studies of non-human primates showed thatfemale skin coloration is influenced by circulating estrogens and thus reflect
reproductive status. This suggests that the link between red color and fertility may haveroots in human biological heritage and emerged in a shared primate ancestor. The
current research project tests the hypothesis that woman lips coloration plays a role into
the human sexual communication as a visual modality providing information about theprobability of ovulation. I predicted that lips become darker and redder (i.e., stronger
signal) during period of high vs. low probability of ovulation. I collected digital
photography (woman face) and salivary samples of 14 women (mean = 25.6 ± 3.4
years-old, 1-2 menstrual cycles per participant, 2 participants using hormonal
contraceptive -controls) at the Primate Research Institute of Kyoto University between
April and July 2018. I collected digital images and saliva samples 3 times across themenstrual cycle, i.e., at Day 5 (follicular phase, low probability of ovulation), Day 14(high probability of ovulation), and D 23 (luteal phase, probability of ovulation). For
each image, I calculated lips luminance (achromatic parameter) and redness (chromaticparameter) under CIE standards. Saliva samples were analyzed for estradiol. I analyzed
the relationship between lips coloration (redness and luminance), cycle phase, andestradiol in R by fitting a series of GLMMs. I did not find evidence for a link between
lips coloration and salivary estradiol. However, I found that lips redness, but notluminance, significantly varied across cycle phases with lips becoming less red during
the luteal phase vs. follicular phase and estimated ovulatory period. Although thesample size of the present study is small, this is the first evidence for a link between
woman lips coloration and reproductive status, suggesting that lips coloration may act
as a sexual signal. The next step of the study will aim at increasing sample size,
resolution (number of data point per menstrual cycle), and men/women perception oflips coloration.

16

O-11

Images speak louder than numbers? Evaluation of liver function by
ultrasound and serum biochemistry in owl monkeys
Rafaela Sayuri Cicalise Takeshita1*; Leandro Nassar Coutinho2; Maria Vivina Barros Monteiro3; Pedro
Mayor4; Paulo Henrique Gomes de Castro5; Sheila Canevese Rahal6; Frederico Ozanan Barros Monteiro2
1 Primate Research Institute,Kyoto Universit, Inuyama, Aichi, Japão, 2 Universidade Federal Rural da Amazônia(UFRA), Belém, Brasil, 3 Universidade Federal do Pará (UFPA), Castanhal, Pará, Brasil, 4Universitat Autònoma deBarcelona, Bellaterra, Barcelona, Espanha, 5Centro Nacional de Primatas (CENP), Ananindeua, Pará, Brasil,
6Universidade Estadual Paulista (UNESP), Botucatu, São Paulo, Brasil*rafaela.takeshita.32c@kyoto-u.jp

Owl monkeys (Aotus azarai infulatus) are the only nocturnal anthropoids, and have been recently the
focus of field and lab experiments to understand what physiological adaptations are distinguishable fromdiurnal primates. For this reason, the maintenance of this species in captivity has increased, as well as theneed to conduct regular health check-ups to monitor their health. The liver is one of the main organs
involved in physiological regulation, due to its functions in biosynthesis, catalysis, detoxification,
digestive, and immunologic. Hematological tests combined with analyses of liver enzymes, such asalkaline phosphatase (ALP), alanine (ALT) and aspartate aminotransferase (AST), bilirubin (BIL), andgamma glutamyl-transferase (GGT) are commonly used in biochemistry exams of the serum to screen forhepatic diseases. However, whether these enzymes can accurately detect liver diseases remains unclear.
The objective of this study was to use ultrasound to evaluate the liver of captive owl monkeys, andcompare with their serum biochemistry exams. The subjects were 69 captive owl monkeys from the
National Primate Center (CENP, Brazil) by ultrasound, serum biochemistry and hematological analyses,
taking in consideration their age, sex, and health condition, assessed by clinical examination, andparasitological tests. The results showed that 16 individuals had parasites and were excluded from theanalyses. Nine individuals had liver disorders and 41 were healthy. The hematological parameters andserum biochemical values were compared between these two groups using Mann-Whitney tests, todetermine how those values are affected by the liver appearance, and checked manually with the reference
values for the species to verify their ability to detect liver disorders. Among the nine individuals affected,
only 2 showed an increase in ALP activity. One of those individuals also showed high count oflymphocytes and monocytes in the hematological exam. Two individuals had elevated count of leukocytes,
eosinophils, lymphocytes and monocytes, and 1 individual showed a slight increase in red blood cells
count, but no change in other parameters. The statistical tests showed that animals with liver disorders
had higher concentration of leukocytes and lymphocytes in comparison with the healthy group, but thethere were no significant differences on biochemical parameters. Biochemical and hematological exams
are useful, but do not accurately detect liver disorders. The combination of imaging diagnostic methods
with those exams can aid in a complete understanding of the liver function and may aid in early detectionof hepatic diseases in owl monkeys.

17

O-12

Sensitivity to Bitter Molecule Phenylthiocarbamide (PTC) in FourSpecies of Sulawesi Macaques

Kanthi Arum Widayati*1,6, Yan Xiaochan2, Nami Suzuki-Hashido3, Akihiro Itoigawa2,Laurentia Henrieta
Permita Sari Purba 1 , Fahri Bajeber4, and Yohey Terai5, Bambang Suryobroto1, Hiroo Imai2
1

Department of Biology, Bogor Agricultural University, Bogor, Indonesia, 2 Primate Research Institute,

Kyoto University, Inuyama, Japan, 3 Academy of Emerging Science, Chubu University, Kasugai, Japan,
4

Department of Biology, Tadulako University, Palu, Indonesia, 5Department of Evolutionary Studies of

Biosystems, The Graduate University for Advanced Studies, Hayama, Japan, 6Wildlife Research Center,
Kyoto University, Kyoto, Japan*

kanthiarum@gmail.com; widayati@ipb.ac.id

Bitter perception plays an important role in avoiding ingestion of bitter-related toxins by
inducing innate avoidance behaviour in mammals. Bitter tasting is mediated by the Gprotein-coupled receptor TAS2Rs, which are located in membrane of the sensory cell. Oneof the best-studied bitter taste molecules is phenylthiocarbamide (PTC) that is recognized
by TAS2R38 receptor. In macaque, experimental study of avoidance behaviour of PTC and
its in vitro functional assay were done only in Macaca fuscata. Most M. fuscata individuals
are sensitive to PTC, while the non-sensitives are found in restricted area that is interpreted
as an adaptation to particular environments. It might be speculated that specific geography
allows independent divergence of bitter perception. Seven species of Sulawesi Macaques
distribute allopatrically with restricted parapatry therefore they might be hypothesized as
evolving different bitter perceptions. In this study, we did experimental behavior andgenetic characterization of TAS2R38 of M. tonkeana, M. hecki, M. nigrescens and M. nigra.
We would like to know if there are variations in bitter tasting and its genetic basis of thefour species. We tested avoidance behaviors of 54 individuals of M. hecki (N: 16), M.
tonkeana (N: 12), M. nigra (N: 15) and M. nigrecesense (N: 11). We found all M. hecki
individuals avoided apple pieces soaked in PTC solution indicating that they are
PTC-sensitive, while in other three species not all of them are PTC-sensitive; those are,eight individuals of M. tonkeana, 13 individuals of M. nigra, and 10 individual of M.
nigrecense rejected the PTC. These results showed that there are polymorphisms in bitter
perceptions of three species. Molecular genetic analyses reveal 20 TAS2R38 haplotypes
with different translated amino acid sequences from all four species. There is one shared
PTC-sensitive haplotype among the four species. We confirmed the TAS2R38 phenotype of
this PTC sensitive using functional analysis. The half maximal effective
concentration (EC50) of the PTC receptor is 1.93 mM, while that of the PTC non-sensitive
cannot be determined. We found that the genetic bases of PTC non-sensitive phenotype aredifferent in each species. In M. tonkeana, relative to the PTC-sensitive phenotype, thenon-sensitive showed amino acid substitution on position 117, 130, 134. In M. nigra, thePTC non-sensitive phenotype showed a premature stop codon at amino acid position 178.
We have not yet analyzed the cause of PTC non-sensitive phenotype in M. nigrecense.
These results indicate that the TAS2R38 receptors evolved independently in each species,
however whether these were the results of adaptation need to be studied.

18

O-13

How to rejuvenate the super-aged society

Kaoru Sekiyama1*
1

Graduate School of Advanced Integrated Studies in Human Survivability, Kyoto University, Kyoto,Japan*sekiyama.kaoru.8a@kyoto-u.ac.jp

Japanese people are expected to live until about 90 years old, while most of themshow gradual increase of needs for assistance in daily living from around 75 years. A
major cause of such needs is dementia that more than 40% of the population suffer by
age 85. Although Japan is the most advanced country in population aging, global aging
is an important issue. My recent research has focused on questions about what lifestyles
are appropriate to rejuvenate older adults’ brain in normal aging. I will show some ofour studies on effects of physical exercise as well as musical instrument practice.

19

O-14

Working for International Organizations

Eriko Ishida Kawai1
1Graduate School of Advanced Integrated Studies in Human Survivability, Kyoto University, Kyoto, Japan

Working at an international organization (IO) has both positive and negative aspects.
Positive aspects include diversity in terms of nationality and gender, excellent
compensation packages, good work life balance, and a multicultural environment. As
international organizations promote social justice and equity, in general they provide
better career prospects for women than in Japan.
The negative aspects include lack of funds and favoritism such as political
appointment. As the organization does not grow in size, it is often difficult to get a
promotion. International bureaucracy and internal politics induce frustration andinefficiency. Organizational issues include turf wars because of an overlap of themandates. For example, collaboration among WHO, UNICEFF, UNPF in the field of
medical care, and public health is dysfunctional. This leads to significant criticism
concerning the bureaucracy and lack of accountability.
For those who are interested in working at IO, languages skills (English plus one
official language) and higher education of at least MA and preferably PhD level are amust. One should be able to write documents fast and need good diplomatic andcommutation skills. In addition, work experience in related fields are often necessary.
As Japanese nationals are under-represented, the Japanese government provides special
recruitment opportunities. Internships are a very good way to get a position by proactive
networking.

20

O-15

Working for an international organization: communication skill

Yoshihiro Kawai1,2
1Graduate School of Management, Kyoto University, Kyoto, Japan2

Wildlife Research Center, Kyoto University, Kyoto, Japan

Essential skill in working for an international organization is communication skill.
Communication skill is not the same as language skill. The presentation will provide
key elements for effective communication in an international organisation and will
describe practical ways to improve communication skill.

21

O-16

Filling the gap: an update from the Kyoto City Zoo and an invitation
to the ICEE2019

Yumi Yamanashi12*, Yoko Sakuraba1,2, Hironori Torii1,3, Fumio Itoh1 and Nobuaki Iwahashi1
1
2

Kyoto City Zoo, Kyoto, Japan

Wildlife Research Center, Kyoto University
3*

Faculty of Science, Kyoto University

yumi.yamanashi.kycz@gmail.com

Ensuring good health and well-being of animals is an essential part of modern zoo
management. However, it is often the case that some charismatic animal species attract
more attention than others. As a result, variations in the level of environmental
enrichment across different species occur, irrespective of the actual needs. In order tofill in the gap at the Kyoto City Zoo, we attempted gap analysis and environmental
enrichment based on the analysis. For the first step, keepers rated animal enclosures ofwhich they are in charge, using welfare assessment sheets developed by a NPO Wild
Welfare. After assessing the 51 enclosures, we discussed and decided the target species
to pay more attention for the year. The first prioritized species were scarlet macaws andgreat green macaws. We conducted environmental enrichment and assessed its effect,collaborating with intern students. We will discuss the potential values on this approachfrom the view of animal welfare and education. In addition to the case study from thezoo, we will also introduce the plan of ICEE 2019.

22

O-17

Verification of the effectiveness of environmental enrichment
and the influence by visitors in captive tigers

Momoko Oka1*, Yumi Yamanashi2, Kota Okabe2, Masayuki Matunaga2 and Satoshi Hirata1
1

Wildlife Research Center, Kyoto University, Kyoto, Japan
2

Kyoto city Zoo, Kyoto, Japan*

momo950322@yahoo.co.jp

Zoo animals are often bred in environments that are significantly different from their
original habitats. As a result, abnormal behaviors such as pacing around in the exhibit,
shaking the body in front and back, left and right, coprophagy and regurgitation may
occur. Among them, pacing is common in large felid species such as tigers.
Environmental enrichment is a variety of devices for improving the welfare of zoo
animals and is considered important for decreasing pacing. Several factors such as thepresence or absence of enrichment and the influence of visitors are involved in thepacing, but such factors were rarely investigated simultaneously. In this study, weinvestigated the effectiveness of enrichment and the factors influencing the behavior of
captive Amur tigers (Panthera tigris altaica).
We observed the behavior of 3 amur tigers in the Kyoto city Zoo and examined theeffect of types of enrichment, temperature and number of visitors on behavior. The
result indicates that there are individual differences in the preference of enrichment.
Moreover, as the maximum temperature gets higher, the pacing frequency decreases andthe resting frequency tends to increase. The influence of the visitor effect is not clearlyseen as much as that of the temperature. It can be said that temperature is more
important as a factor influencing the behavior of the tiger. Since there are individual
differences in the preference of enrichment, it is considered that performing multiple
enrichment is useful for tigers, and is helpful in grasping individual preferences.

23

O-18

Salivary alpha-amylase enzyme as a non-invasive biomarker of acutestress in Japanese macaques (Macaca fuscata)1
1

Nelson Broche Jr.

Primate Research Institute, Kyoto University, Inuyama, Aichi
broche.nelson.63a@st.kyoto-u.ac.jp

Human studies have revealed salivary alpha-amylase (sAA) enzyme levels are
positively correlated with increases of the hormone norepinephrine, allowing sAA to act
as a biomarker for sympathetic nervous system (SNS) activity. The SNS is associatedwith the fight-or-flight response and is a separate but parallel stress response system tothe hypothalamic-pituitary-adrenal (HPA) axis. Recent non-human primate studies havebegun investigating the potential use of sAA as a stress biomarker in rhesus macaques,
gorillas, and other species. Validation of sAA enzyme as an acute stress biomarker inJapanese macaques could provide a useful tool for future research questions as well aspractical uses in animal welfare. However, there are currently no published reports ofsAA validating its use as an acute stress response marker for Japanese macaques. Saliva
collection through non-invasive means from unrestrained monkeys can be a challenge
and impede such work. Developing a non-invasive method for cooperative saliva
collection between the researcher and monkey is not only necessary for accurate datacollection but also ethically sound. This study had a two-fold aim: [1] develop asystematic method for non-invasive saliva collection by creating a cooperativeenvironment with the study subject and [2] validate sAA as a biomarker of acute stressin M. fuscata. I will discuss developing a methodology for cooperative saliva collection,
results of 22 stress tests, limitations, and future directions.

24

O-19

“Bar hanging” behavior: a look into a potentially culturally propagatedbehavior and its relation to stress

JS Alejandro1*, H Monier2, F Bercovitch1,3,4 and MA Huffman1
1

Primate Research Institute, Kyoto University, Inuyama, Aichi, Japan2

École Nationale Vétérinaire de Toulouse, Toulouse, France
3

Save the Giraffes, San Antonio, TX, USA

4

Wildlife Research Center, Kyoto University, Kyoto, Japan*josue.pastrana.64s@st.kyoto-u.ac.jp

In recent decades, what was thought to be a unique human trait- culture-, has beenchallenged and well demonstrated to exist in various non-human primate species. Oneof the Japanese macaque groups housed at the Primate Research Institute of Kyoto
University engages in a unique behavior that is not seen in any of the other captive
groups, known as “bar hanging”. The animals are in good health and are kept in
identical conditions as many other groups, yet many animals practice this behavior
exclusively in this group. Some observers see this as a play behavior while others see it
as a stereotypic behavior related to stress or a way of coping with stress. With datacollected over the past 15 years in the form of videos, behavioral data, genetic data,physiological data, we attempted to elucidate 1) how this behavior might have spreadand learned over time, 2) what is its relation to stress, if any, 3) understand thedemographics of individuals who engage in this behavior (age, sex, rank, kinship). Our
predictions were that animals that were more closely related to each other would show
similar patterns of the behavior and animals who engage more frequently in thisbehavior had lower levels of faecal glucocorticoids. We believe our work can contributeto the increasing literature on primate traditions and how monitoring these behaviors
can be an important tool in behavioral management.

25

O-20

Studies on ecology and behavior for wildlife conservation
Shiro Kohshima1*
1

Wildlife Research Center, Kyoto University, Kyoto, Japan* kohshima@wrc.kyoto-u.ac.jp

We study the ecology and behavior of various animals, especially large animals that have not been wellstudied, because we believe that understanding the ecology and behavior of each animal species isessential for its conservation and the mitigation of its conflicts with humans. We focus on large animals
because many of them are endangered species, umbrella species and flagship species, important for the
ecosystem-scale conservation of their habitats. For example, Borneo orangutans and Amazon riverdolphins can symbolize ecosystem conservation of Asian tropical rainforests and the Amazon River,respectively, and their conservation can contribute to the conservation of many other organisms. We havestudied the ecology and behavior of Asian elephants, Malayan tapirs, orangutans, wolves, wild dogs,
civets, porcupines, sloths, otters, sea otters, Amazon River dolphins, Ganges River dolphins, and Amazon
manatees, as well as various dolphin species, including killer whales and seal species. We study theseanimals in the wild and/or captive condition. We study wild animals using various advanced methods
such as bio-logging, camera trapping and sound/image analysis. Precise analyses of the behaviors
observed in captive environments, such as zoos or aquariums, are also important to understand thefunction and meaning of each behavior. So, we officially collaborate with 11 zoos and 7 aquariums inJapan to promote studies in captivity. We collaborate with these zoos and aquariums to improve thecaptive condition of the animals and their quality of life, based on the scientific results.
We organize a JSPS core-to-core program “Center of Excellence for Conservation of Tropical
Bio-diversity focusing on Large Animal Studies”, which aims to promote international academic
exchange for tropical bio-diversity conservation among Japan, India, Brazil, Malaysia, China, Indonesia
and the UK. In this program, we promote collaborative studies on various large animals in each member
country and conduct international training courses in Japan. We also organize international seminars
and workshops to realize the “Field Museum”, a new generation zoo/aquarium/botanical garden in thenatural habitat of wildlife, which can contribute to research, conservation, education and ecotourism ineach member country. Field Museum is a network of nature reserves and observation facilities in thenatural habitats of wildlife, where we can observe various organisms in captive, semi-captive and wildconditions, as well as conduct research and conservation activities. The Field Museum can contribute not
only to study, conservation, and environmental education, but also to the local community throughecotourism and as a center for conservation of local ecosystems. To realize a Field museum in the
Amazon, we organized a JST/JICA SATREPS Program: Biodiversity Conservation in Amazon, based on
the new concept of the “Field Museum” in collaboration with the National Institute for Amazon Research(INPA, Manaus, Brazil).

26

O-21

Eavesdropping on elephants: How and what do Asian elephants talkwith each other?

Nachiketha Sharma1, 2*, Shiro Kohshima1, Raman Sukumar2,3
1
2

Wildlife Research Center, Kyoto University, Kyoto, Japan

Center for Ecological Sciences, Indian Institute of Science, Bengaluru, India
3

Institute for Advanced Study, Kyoto University, Kyoto, Japan*nachi@wrc.kyoto-u.ac.jp

Communication plays a vital role in the life of all social animals. According to theirevolutionary and ecological needs, animals have evolved various communication
systems such as acoustic, chemical, visual, tactile etc. These systems have far-reachingeffects on social relationships, reproduction and survival of a species which enables us
to understand the biological processes from proximate to ultimate level. Even though,animals communicate with each other by using different forms of communication, here
we concentrate mainly on acoustic signals by using free-ranging Asian elephants as amodel system. Unlike Asian elephants, the vocal communication in African elephants
has been well studied. African elephants are known to have complex vocal networking,and well developed referential alarm calls. Therefore, we make an attempt to answer a
few fundamental questions such as vocal repertoire, calling rates, communication withina group, use of alarm calls in Asian elephants which live in fission-fusion society. This
study was conducted in Bandipur and Mudumalai Tiger Reserve (India) which is a partof Nilgiri Biosphere Reserve and harbours one of largest wild Asian elephants’population in the world. We conducted systematic surveys and followed elephants onfoot to record their vocalizations. We quantified vocalizations and associated behaviourof free-ranging Asian elephants during social interactions (intra and intergroup), humandisturbance (humans on foot, vehicular disturbances) and hetero-specific species (birds,
ungulates, predators) within the protected area. We found that Asian elephants mainly
used four call types (trumpets, rumbles, chirps and roars) with slight variants dependingon the context. Out of four call types, roars seem to elicit physical interactions whereas
chirps had no responses from conspecifics. Similarly, fundamental frequencies (F0) of
rumbles produced during disturbance had more modulations compared to social rumbles.
Understanding these systems not just provides detailed information on life history traitsof a species, it also helps in formulating effective conservation strategies in the
Anthropocene epoch.

27

O-22

Short-term gut microorganism dynamics induced by gumnivorous
feeding for lesser slow loris

Kei Matsushima1, Yumi Yamanashi1,2, Fumihiko Okumura3, Mari Hirosawa1,3, Yui Fujimori3,
Yumiko Terao3, Ryo Sato4, Masayuki Nishino4, Sayaka Tsuchida5, Kazunari Ushida5,
Takashi Hayakawa3,61

Wildlife Research Center of Kyoto university, Kyoto, Japan, 2 Center for Research and Education of
Wildlife, Kyoto City Zoo, Kyoto, Japan, 3Japan Monkey Centre, Aichi, Japan, 4 Chubu University
Academy of Emerging Sciences, Kyoto, Japan, 5 San-Ei Gen F.F.I.,Inc., Japan, 6 Primate ResearchInstitute, Kyoto University, Aichi, Japan*

kmatsu@wrc.kyoto-u.ac.jp

Lessor slow lorises (Nycticebus pygmaeus) naturally ingest plant gum as a main food.
However, it is not popular to feed plant gum for slow lorises in captive environments
such as zoos. In this study, we fed plant gum continuously for two lesser slow lorises
reared in Japan Monkey Centre, which have rarely eaten plant gum. We assess the effectby analyzing the change of their gut microbiota from the viewpoint of animal welfare
and feeding enrichment. Plant gum is composed of low degradable polysaccharide.
Therefore, slow lorises may depend on gut microbiota for digestion of plant gum.
We collected their feces before and after feeding gum arabic for one month,respectively. We amplified the V1-V2 region of the bacterial 16S rRNA using universal
primers by PCR and massively sequenced by an Illumina Miseq. Sequenced data isanalyzed by the Qiime and R platform. We found that the microbiota structure is quitechanged drastically after two days from starting feeding gum, which is the same as theirdetention time. Especially, the most drastic changed happened in Prevotellaceae. Sometaxa had decreased or complementarily increased and some had vanished. We will
analyze the functional aspect of these taxa by cultivation.

28

O-23

Marching with Penguins

Lloyd Spencer Davis
University of Otago, Dunedin, New Zealand

The documentary March of the Penguins (2005) helped perpetuate many
misconceptions about the breeding behavior of penguins that Professor Lloyd Spencer
Davis has spent his career debunking. In contrast to the tagline of the film, he has
shown that in the harshest place on Earth, it is not really love that finds a way. However,it has become apparent recently that Davis has been following in someone else’s
footsteps: those of George Murray Levick.
Here, Davis reports on the breeding behavior of penguins and his current focus on
Levick.

29

O-24

Just like clockwork? On the significance of periodic penguinsAndrew J. J. MacIntosh1, Xavier Meyer1,2, Andre Chiaradia3, Akiko Kato4, Yan Ropert-Coudert4
1

Primate Research Institute, Kyoto University

2

Department of Ecology, Physiology and Ethology, Strasbourg University
3

Research Department, Phillip Island Nature Parks
4

Center for Biological Studies in Chizé

Understanding what drives variation in behavior across individual animals andhabitats is fundamental to ecological research. It is also critical for our ability to predict
how threatened species might cope with environmental change. Working withorganisms that can act as indicator species for environmental change can teach us a lot
about behavior-environment interactions and how they may change over time. In thisrespect, penguins, upper-level predators in the southern marine ecosystem, are excellent
subjects to study. We have been investigating emergent patterns of behaviouralorganization – in this case fractal time – in penguins and a range of other animal taxa tohelp us understand the causes and potential consequences of variation in behavioral
complexity. Data suggest that complexity in temporal patterns of behavior isbiologically adaptive, and that complexity loss, i.e. the greater periodicity or stereotypy
often observed in stressed or diseased individuals, may reflect reduced behavioural
fitness. In this talk, we explore physiological, ecological and phylogenetic determinantsof so-called ‘complexity signatures’ and how they can be used to comment on optimality
in behavior in a given environment. Our ultimate aim is to develop a behavioural index
that will allow us to reliably identify matches and mismatches in animal-environment
interactions, and to thereby help us predict the outcomes of a changing planet on animal
populations globally.

30

O-25

Jelly in the Belly: Why do penguins eat jellyfish?

Jean Baptiste Thiebot1*, Julie C. McInnes2
1

BioScience Group, National Institute of Polar Research, Tachikawa, Tokyo, Japan
2

Australian Antarctic Division, Kingston, Tasmania, Australia*

jbthiebot@gmail.com

There is growing evidence that gelatinous zooplankton (“jellies”) are a regular prey
for a wide array of marine endotherms, including penguins. This interaction has been
documented across the world’s oceans and using varied techniques. Consumption ofjellies is intriguing in terms of energy reward, given endotherms have relatively highenergy demands and jellies would provide little energy as prey. Here we show thatmany of the intuitive hypotheses commonly raised to explain predation on jellies havealready been tested and most of them are unsupported. We emphasize that jellies are
generally a non-anomalous prey (i.e., not reflecting an ecosystem perturbation), andpropose that they might be beneficial to marine endotherms for non-energetic reasons,such as enhancing physiological processes. Changing the “junk-food” hypothesis for a“green tea” framework may thus allow to better understand the widespread predation onjellies, although further biochemical analyses are needed to clarify this point.

31

O-26

Solving the mystery of female-biased stranding in Magellanic penguins

Takashi Yamamoto1*, Ken Yoda2, Gabriela S. Blanco3 and Flavio Quintana3
1

Research and Development Center for Data Assimilation, The Institute of Statistical Mathematics,

Tachikawa, Tokyo, Japan, 2 Graduate School of Environmental Studies, Nagoya University, Nagoya,
Aichi, Japan, 3 Instituto de Biología de Organismos Marinos, Puerto Madryn, Chubut, Argentina* ytaka615@gmail.com

Thousands of Magellanic penguins (Spheniscus magellanicus) strand every austral winteralong the coast of Brazil; this is more than a thousand kilometers away from their northernmost
breeding colony in northern Patagonia. Curiously, females usually outnumber males at strandingsites. Previous studies suggest that penguins move north from their breeding colony to winteroff the coasts of northern Argentina, Uruguay, and southern Brazil, and assume that females
may remain in areas closer to the coast and/or travel farther north than males during thenon-breeding period. However, so far no remarkable differences have been reported in theirmigratory movements between sexes, although tracking data were very limited for early stagesof winter (mostly up to early June), failing to record their at-sea distributions during the peakstranding period (late July–early August); therefore, a mechanism behind the female-biased
stranding in wintering penguins has been an unresolved mystery despite the growing importancefor their conservation.
To examine the different spatial utilizations adopted by males and females, we recorded thewhereabouts of Magellanic penguins throughout the non-breeding period using animal-borne
data loggers (geolocators), which record ambient light intensities and elapsed time, providing
daily locations (latitudes are derived from day lengths and longitudes from the relative timing ofrecorded midday or midnight). Penguins initially migrated northwards as previously reported.
However, females reached more distant wintering areas than males and did not dive as deep,
suggesting that females might avoid vertical competition for food resources (i.e. niche
partitioning), which may be a consequence of the avoidance of intraspecific competition due tosexual dimorphism in body size. Alternatively, thermal constraints may drive sexual differencesin the spatial extent as lower water temperatures induce greater heat loss, resulting in elevatedmetabolic rate. Consequently, females are more likely to suffer environmental hazards along thecoast of Brazil, one of the most threatened marine environments in South America. Our study
highlights the size dimorphism-related differences in spatial usage by males and females during
the non-breeding period, which is probably associated with female-biased greater risk ofmortality through exposure to a wider range of potential threats, which has the critical
implications for effective conservation practices.

32

O-27

Is a Bonobo Older Brother or just Meat for Bongando Ethnic Group?
-Focusing on Local Perceptions towards Bonobos in DRCongo-

Aya YOKOTSUKA1*
1

Graduate school of Asian and African Area Studies, Kyoto University, Japan*aya.yokotsuka@gmail.com

Bushmeat hunting is one of the biggest threats to the great apes in Central Africa. The
Bongando ethnic group in Democratic Republic of Congo traditionally has a taboo
against eating bonobos (Pan paniscus). The reason why Bongando avoid eating bonobo
is because they believe bonobos are from same ancestors with Bongando. Also bonobos
have a no tail, so bonobos are not animals but humans in Bongando’s animal taxonomy.
However, researching other villages, the situation revolving around bonobos isdramatically different. This study aims to examine how local perceptions towards
bonobos become diversified within and beyond the reserve.
I conducted a survey questionnaire to 797 Bongando people living in 7 villages toinquire their human-bonobo interactions. Two of the villages surveyed are from thereserve and five are from the non-protected areas. Results show that the perception
towards the bonobos differed between the people from the reserve and the non-protectedareas. In the reserve, bonobo research has been continued over 40 years.
People from the reserve tend to avoid eating bonobo due to their traditional beliefs
and benefit from the bonobo research. The support from the bonobo researchencourages the preservation of the bonobos in the reserve. Conversely, people from thenon-protected areas are inclined to eat bonobo. This can be seen especially in the agegroup between 20 and 40. Furthermore, the availability of bonobo meat in these areas
tends to be high in the villages from the non-protected areas.
According to my interviews, villagers hunt bonobos not for only sale.
This mean is villagers who hunt bonobos, and also consume bonobo meat bythemself.
It is assumed that cultural interchanges have been accelerated after civil wars.
Plantations around villages were collapsed, so people go to cities to get work.
The traditional taboo of bonobo is disappearing rapidly and the situation is critical in
the non-protected areas.

33

O-28

Social and Sexual Development up to Natal Dispersal in Female
Bonobos

Kazuya Toda1*1Primate Research Institute, Kyoto University, Aichi, Japan*

toda.kazuya.78x@st.kyoto-u.ac.jp

Dispersal from the natal group is of great decision of individuals in their life
histories of social animals, because its farewell to familiar local areas and groupmembers is accompanied with fitness risks. Bonobos are one of few animals to form
male-philopatry groups, but it has remained room to study biological factors of female
dispersal. In my Ph.D. study, I have investigated behavioral and hormonal changes
prior and posterior dispersal in female bonobos to examine proximate causes of female
dispersal. Subjects of this presentation were 7 natal females of two habituatedunit-groups at Wamba, Luo Scientific Reserve, in DR Congo. In total 23 months over
5 years, behavioral observations were collected with ad-libitum and focal animal
sampling during the party following for 3,295 hours, and hormonal analyses wereconducted by Enzyme Immunometric Assay with urine samples. This study foundsocial and sexual changes in female bonobos up to the timing of dispersal (6.9 years
old, N = 7). At puberty, female bonobos would become to prefer senior adult females
while independent of their mothers.

34

O-29

Did life in trees drive the adaptive evolution of taste in koala andprimates?

Takashi Hayakawa1*
1

Primate Research Institute, Kyoto University, Inuyama, Aichi, Japan*

hayakawa.takashi.3a@kyoto-u.ac.jp

Grasping hands with opposable digit is the most distinguishing feature of arboreal
adaptation in primates. Life in trees increased body size in anthropoid primates owing toincreased nutritional supply from woody plants, but they encountered high level of plant
toxins (plant secondary metabolites, PSMs). The evolutionary struggle with plants’
PSMs might have caused the expansion of bitter taste receptor genes (TAS2Rs) inanthropoid primates, which avoid ingesting possibly toxic PSMs. Thus, anthropoid
primates, including human, became one of the groups with the highest numbers of
TAS2R genes among mammals. Changing perspective from the northern to southernhemisphere, there are various species of marsupials, which have different history ofadaptive radiation to various environments from placental mammals like primates.
Fieldwork in Australian forests enable us to find very attractive arboreal marsupial
folivore, koala (Phascolarctos cinereus). Koalas grasp branches of trees like primatesand intake highly toxic eucalyptus leaves as major items of the diet repertoire. Koala
Genome Consortium recently determined whole genome sequences of koalas. Koala
and other marsupial genomes provided information about the taste receptor gene
repertoire. As a result, the koala has the largest number of TAS2R genes, 24, among
analyzed Australian marsupials, which is also the largest level among all mammals.
This is presumably a result of koala-specific gene expansion and evolutionary
convergence with plant-eating primates. Closely-related herbivorous species of koala
are terrestrial such as wombats and wallabies. More comparative genomics ofmarsupials and placental mammals will provide insight into relationship between
arboreality and taste receptor evolution.

35

O-30

Chimpanzees’ behaviour in the prosocial choice paradigm
depends on the position of the token in the feeder

Maciej Trojan1*, Julia Sikorska1,2 and Anna Jakucińska3
1

Department of Ethology and Comparative Psychology, Nicolaus Copernicus University, Toruń, Poland
2

Department of Animal Behaviour, University of Warsaw, Warsaw, Poland
3

Warsaw Municipal Zoological Garden, Warsaw, Poland*

maciej.trojan@umk.pl

Chimpanzees evince prosocial behaviour in nature, as well as in the laboratory research.
In our study we were looking for factors modifying this kind of behaviour using the
prosocial choice paradigm. The results show that many individuals tend to help others,
share resources, and bring protection. Altruism can be examined in a prosocial choiceparadigm. The essence of the research in this paradigm is the ability to provide a benefit toanother individual, with little or no effort from the individual under examination. The aimof this study was to verify whether chimpanzees evince a prosocial behavior and whether
they use the tit-for-tat strategy during token exchaning. During the prosocial paradigm tasks,
each chimpanzee had access to 10 tokens (5 green – prosocial choice - two chimpanzees got
a reward; 5 red - selfish choice - only the chimpanzee which made a choice got a reward).
Tokens were available during the session each day for 10 days. In the first experimentalgroup tokens were mixed after each choice of the individual. Chimpanzees were exchanging
tokens in the one by one system - 20 tokens per day during one session. In the second
experimental group they also gave 20 tokens in the system one by one, but the experimenterput all the tokens in two separate piles - red and green =. Red tokens were always on the left
side, and green tokens on the left side, and vice versa (Depending on the day of theexperiment). In the the control group, one individual chose 20 tokens (mixed) in a row. The
dominance of a particular hand in a specific type of social choice task strongly influences
the color of token chosen by the individual. Because of the tray structure and body position,
chimpanzees chose tokens with the dominant hand on the diagonal. Day by day this effectweakened, but each individual gave tokens which were places diagonally to the dominanthand. The effect was stronger in males because they had a tendency not to change theposition of the body during the session. Females were more active, which resulted in aslightly weaker effect of the impact of motor lateralization. To sum up. In this type ofprocedure we should have a lot of tokens, and every time before making the choice theyshould be well mixed.

36

O-31

Male Reproductive Success in Bornean Orangutans (Pongo pygmaeus)

Tomoyuki Tajima1*, Titol P. Malim2, Eiji Inoue3
1

Lab. of Human Evolution Studies, Graduate School of Science, Kyoto University, Kyoto, Japan
2
3

Sabah Wildlife Department, Kota Kinabalu, Sabah, Malaysia

Department of Biology, Faculty of Science, Toho University, Chiba, Japan*tajima002@gmail.com

Among primates, male reproductive success is not always associated with dominancestatus. Male orangutans exhibit intra-sexual dimorphism, and clear dominancerelationships exist among the males. Previous studies reported that both the morphs
were able to sire offspring but the information is still limited. The present study aimed
to investigate the difference in reproductive success between the two male morphs in
Bornean orangutans. We conducted sample collection and behavioral observation in afree-ranging population of Bornean orangutans. We assigned the paternity of eight
infants using 12 loci of microsatellite. A flanged male sired most of the offspring bornfrom parous females, and an unflanged male sired a firstborn offspring. This isconsistent with behavioral observation: the dominant flanged male showed little interest
with nulliparous females, in contrast to the unflanged males, which frequently matedwith them. This suggests that the dominant flanged male might be able to monopolize
most of the fertilizations of parous females, while unflanged males might seek any
mating opportunity available in the absence of the flanged male and copulate withfemales, including nulliparous ones, even though their conception probability isrelatively low. We are conducting a paternity analysis in Danum Valley, targeting on thewild population of Bornean orangutans.

37

O-32

Research, Conservation and Management of the Proboscis Monkey
Populations in Klias Peninsula, Sabah, Malaysia

Henry Bernard1*
1

Institute for Tropical Biology and Conservation, University Malaysia Sabah, Sabah, Malaysia*

hbtiandun@gmail.com

The proboscis monkey, Nasalis larvatus; a Bornean endemic species, is one of the
major eco-tourist attractions on the island of Borneo, including in the Malaysian Stateof Sabah located in northern Borneo. Despite its economic importance, much of theproboscis monkey’s key habitats, i.e. riparian and coastal mangrove forests, have been
greatly reduced or disturbed and fragmented as a result of logging activities,
infrastructural and agricultural development, human settlements expansions, and forest
fires. The population size of proboscis monkey in Sabah was estimated to be close to
6,000 individuals, with five major centres of continuous populations and numerous
small isolated sub-populations. The Klias proboscis monkey population is one of thestrongholds of proboscis monkey populations in Sabah with estimated population size
ranging between 400-600 individuals. It is also regarded as the only viable populationof this monkey on the west coast of Sabah. Since 2005, the Institute for Tropical
Biology and Conservation, and its partners, have conducted several studies on theproboscis monkey in Klias. This paper presents the main findings of the research withthe main aim to contribute knowledge that are useful for the conservation management
and monitoring of the proboscis monkey population in Klias.

38

O-33

The way forward of wildlife research

Miho Murayama1*
1

Wildlife Research center, Kyoto University, Kyoto, Japan*

mmurayama@wrc.kyoto-u.ac.jp

The Wildlife Research Center of Kyoto University was established as the first
university wildlife research institution in Japan in April 2008. In the past decade, wehave expanded this network within the country and abroad, conducting research andeducation. Currently, we collaborate with more than 20 zoos and aquariums to conductex-situ conservation. We conduct in-situ conservation research in cooperation with local
research institutes at seven overseas research bases. We accept graduate students of the
Graduate School of Science to accomplish our education mission. Also, we participate
in the leading graduate program, “Primatology Wildlife Science (PWS)”. So far morethan 60 students have studied more than 38 species of wild animals in 13 countries.
Although we are still a small organization, we have created the new discipline, “wildlife
science”. At its inception, the center started with six full-time faculty members. Now,
36 professors work on special posts and concurrent posts. This development would nothave been accomplished without the dedicated cooperation of the professors andadministrative staff of the institute.
In my research field we are applying molecular analysis to learn more about wild
animals. We collected DNA samples of over 27,000 individuals representing 200
species of mammals noninvasively from feces and hair. Using this DNA Zoo, we are
identifying subspecies/populations/kinships/sex by genotyping polymorphic markers.
We are also surveying individual differences in functional genes related to personality
traits (e.g. stress susceptibility) and reproduction. As a new approach we are trying toestimate age by epigenome analysis.
We celebrated 10th anniversary in 11th June. In the next decade, we will pursue evengreater possibilities in education and research, together with young researchers who
have advanced to realize our mission of wildlife conservation. Again, I would like toexpress my gratitude to all participants for your support, now and in the future.

39

O-34

The Arctic Tundra & the African Savannah: Common Conservation
Conundrums Confronting Caribou and Giraffes

Fred B. Bercovitch1, 2, 3*
1
2
3

Save the Giraffes, San Antonio, TX, USA

Wildlife Research Center, Kyoto University, Kyoto, Japan

Department of Animal, Wildlife, & Grassland Sciences, University of the Free State, Bloemfontein,South Africa*

fred.bercovitch@savethegiraffes.org

Although giraffes and caribou occupy quite different habitats, they face similarconservation challenges. In this presentation, the life history of both types of animals
is summarized, followed by noting the major conservation issues that confront the two
species. The IUCN Red List status of giraffes and caribous is presented and compared.
Some of the scientific and political problems inherent in listing a species as threatenedwith extinction are discussed. Finally, the prospects for future conservation aresummarized, along with noting a fundamental disagreement among conservation
biologists as to what is the most productive tactic to adopt for saving species.

40

O-35

Culture, social learning and death in Japanese monkeys

C F I Watson1*, T Matsuzawa2
1
2

Wildlife Research Center, Kyoto University, Kyoto, Japan

Institute for Advanced Study, Kyoto University, Kyoto, Japan*

cfi.watson@gmail.com

In this talk I will give an overview of my research on Japanese macaques. I havecarried out a literature survey of potential cultural variants in this species. Transmission
studies in socially housed captive Japanese macaques have allowed me to assess ability
to socially learn cultural variants. During my research on culture, I took the opportunity
to observe responses of Japanese macaque mothers to their dead infants and this led to apaper outlining a framework for future, systematic, quantitative research on this topic.

41

O-36

Bats: How to study animals you cannot see

David A. Hill 1*
1

Wildlife Research Center, Kyoto University, Japan* hill.david.2u@kyoto-u.ac.jp

More than 1,300 species of bat have been described, which is about 20% of all known
extant mammal species. They can be divided into two groups: around 170 species are
Old World fruit bats that navigate using vision and find their food by sight and smell;
the remaining 1,130 or so species (collectively known as “microbats”) all use ultrasonicecholocation to obtain information about their environment. Microbats are difficult
animals to survey and study because they are small-bodied, fast-moving and most active
at night. Also, although they are very vocal, most of their vocalisations are ultrasonic,
so we cannot hear them. At the same time microbats are an important group to study forvarious reasons, both ecological and economic.
There are strong indications that bat populations around the world are being lost or
severely depleted because of habitat loss and degradation, and other forms of humandisturbance. However, because they are difficult animals to survey, it is usually
impossible to estimate the scale of the decline. For the vast majority of species there areno historical data on population size, or even usable estimates of current populations.
I will briefly outline the pressures that microbats currently face and some of themethods that have been developed to survey and study them. I will focus on work I havebeen involved in that uses acoustic lures to significantly improve capture success, andtherefore survey capability. I will illustrate the use of this technique with a few
examples of recent applications.

42

O-37

Evolution of the loss of olfaction in whales provides new perspectives
on conservation of the marine amniotes

Takushi Kishida1*
1

Wildlife Research Center, Kyoto University, Kyoto, Japan*kishida@wrc.kyoto-u.ac.jp

Transition from land to water is one of the most striking types of the evolutionary
events in the history of life. It has been suggested that the olfactory receptor (OR) genes
are functionally divided into two groups: OR genes for detecting water-soluble
molecules and that for detecting airborne molecules. Thus, the OR gene repertoires will
generally be changed profoundly upon aquatic adaptation. Cetaceans are an order ofmammals derived from terrestrial artiodactyls, and all extant cetaceans are fully aquatic.
In this study, I annotated the OR gene repertoires from genome assemblies of all
cetaceans available in public databases and a hippopotamus, and analyzed theevolutionary gain and loss of OR genes during the evolution of cetaceans. Cetaceans
migrated from land to water in the Eocene epoch, and the OR gene repertoires ofamphibious Eocene whales were also inferred from the cribriform plate size of fossil
specimens. Whales are suggested to have lost OR genes required to avoid inedible foods
at this point in evolution. Based on these data, I will discuss about the aquatic
adaptation and the loss of olfactory capabilities among marine amniotes.

43

P-02

Seasonal changes in bats occurrence and activity in the Janówek Fort
(Masovian District, Central Poland)
Judyta Gulatowska1*, Marek Kowalski2, Grzegorz Lesiński3
1

Department of Ethology and Comparative Psychology, Nicolaus Copernicus University, Toruń, Poland
2
Wildlife Society 'Stork', Warsaw, Poland
3
Department of Animal Environment Biology, Warsaw University of Life Sciences, Warsaw, Poland*

jgulatowska@umk.pl

In Central Europe abandoned military objects are often used by bats as winter shelters,
especially in the areas where no natural caves are present. When the temperature outside
falls several degrees below zero perfect conditions for hibernation are created in forts thatare partly under the ground. In the presented study the winter community structure andactivity of bats, occurring in an abandoned fort in Janówek were observed. The aim was
to find out if the fort is used only in winter or whether it serves some purpose also in otherseasons. Number of bats was monitored regularly during the whole year. In the course ofcontrol nights nets were installed in front of entrances to catch and count flying
individuals. Bats staying inside the fortress were also counted. Eight species of bats wererecorded, including relatively numerous occurrence of pond bat Myotis dasycneme. There
were recorded maximum 142 individuals hibernating at the same time, whereas the total
number of bats caught into nets during 20 nights was 341 individuals. The observation ofchanges in bats’ activity near the fort showed that it was used with different intensity but
throughout the whole year. The different dynamics of using the fort by the two the mostnumerous species: Daubenton’s bat and Natterer’s bat was observed. For Daubenton’s
bat Myotis daubentonii Janówek is the place of autumn swarming.

44

P-03

Genetic diversity analysis of the Ryukyu flying fox
Yuto Taki1*, Christian Vincenot2, Yu Sato1, Miho Inoue-Murayama1
1

2

Wildlife Research Center, Kyoto University
Department of Social Informatics, Kyoto University*

sakyoulemmus@gmail.com

There are about 130 mammal species in Japan, and more than 25% of them are bats.
The Ryukyu flying fox is one of the bat species in Japan, distributed in Nansei islands.
They are quite different from small bats. When they spread their wings, the length willreach 80cm. They mainly eat fruits, and sometimes eat flowers and leaves. It is said thatthey have an important role in pollination and seed dispersal. They are divided into 5
subspecies, and 2 of them are regarded as Critically Endangered. The purpose of thisstudy is to evaluate genetic diversity and genetic differentiation of the Ryukyu flying fox
between each island. There are mitochondrial DNA markers which can be used for the
Ryukyu flying fox already. Therefore, we made mitochondrial DNA analysis by using
them. Tissue samples and feces samples were collected in Nansei islands, and DNA
extraction was conducted. We checked mitochondrial haplotypes to compare thembetween the islands.
Key words: genetic diversity, Ryukyu flying fox, mtDNA marker, conservation biology

45

P-05

Do primates yuck?
Pathogen and parasite avoidance behaviors in Papionini and Hominini
Cecile Sarabian1* & Andrew J. J. MacIntosh1
1

Primate Research Institute, Kyoto University, Inuyama, Japan*

sarabiancecile@gmail.com

Parasites and pathogens are ubiquitous in nature and can be costly to animal fitness, sohosts have evolved behavioral counterstrategies to mitigate infection-risk. We conducted
feeding-related
infection-avoidanceexperiments
with5
speciesof Papionini and Hominini: free-ranging Japanese macaques (Macaca fuscata fuscata) of
Koshima Island; captive chimpanzees (Pan troglodytes troglodytes), semi-free ranging
mandrills (Mandrillus sphinx), and group-housed long-tailed macaques (Macaca
fascicularis) at the International Medical Research Centre of Franceville, Gabon; andwith semi-free-ranging bonobos (Pan paniscus) at Lola ya Bonobo Sanctuary, Kinshasa,
Democratic Republic of the Congo. In parallel, we tested whether the behavioral
tendencies observed toward these contaminants correlate with parasite infection. We now
know that hygienic tendencies in Japanese macaques are good predictors of geohelminth
infection (GLMM, all p < 0.01). Then, we expanded our experimental protocol to include
visual, olfactory and tactile cues of feces, blood, SIV+ semen, rotten meat and rotten fruit.
Subjects manifested increased latencies to consumption of food rewards, maintenance ofgreater distances from contaminants, and/or outright refusals to consume food rewards incontaminated versus control conditions (GLMM, all p < 0.1). We are now testing whether
individuals with greater tendencies to avoid potential sources of contamination are lessprone to infection than less sensitive individuals. The avoidance behaviors observed hint
at the origins of hygiene and disgust in humans, and further comparative research is now
needed.

46

P-07

Where do female bonobos migrate? : Implication from mitochondrial
DNA analysis
Shintaro Ishizuka1, Kazuya Toda1 and Takeshi Furuihi1
1Primate Research Institute, Kyoto University, Aichi, Japan*ishizuka.shintaro.37x@st.kyoto-u.ac.jp

Understanding patterns of female dispersal in non-human primates can contribute toelucidate social systems in human ancestors. Bonobos (Pan paniscus), which is one of
the evolutionally closest species to humans, show strong tendency of female dispersal. In
this species, affinitive interactions between females of different groups are often observed
during intergroup encounters. This suggests that females may migrate into neighboringgroups, and mother-daughter dyads might exit between neighboring groups. We
investigated dispersal patterns of female bonobos among three neighboring groups at
Wamba. By analyzing mitochondrial and autosomal genetic markers, we identified
mother –daughter dyads between immigrant females within the three neighboring groups.
We found one mother-daughter dyad within the three groups. This suggests that migration
into neighboring groups is probable in female bonobos. On the other hands, the observed
number of mother-daughter dyad existing within the three groups was not high comparedto the expected number of those dyads in total. Moreover, genetic distance between
neighboring groups tended to be similar with between different populations in same
regional cohorts. These suggest that female immigration into neighboring groups is atleast not frequent in bonobos. This dispersal pattern is considered because female transfer
occurs only when neighboring groups encounter each other.

47

P-08

Ordering strategies of three-choice task by
free-ranging Japanese macaques
Shenwen Xu11*, Kazunori Yamada2, Masayuki Nakamichi2, Masaki Tomonaga1
1Primate Research Institute, Kyoto University, Japan
Graduate School of Human Sciences, Osaka University, Japan2*

vassxu@gmail.com

Behavioral plasticity allows animals to maximize benefits and minimize costs. While
foraging, besides the basic costs of time and energy, group-living individuals are further
exposed to intergroup competition. We investigated how animals adjust behavior to make
optimal decisions in a group-setting foraging task. We conducted a field experiment at aJapanese macaque provisioning site on Awaji island. We designed a three-choice task
where participants came voluntarily and had the option to pull weighted boxes baited withfood in any order. The three boxes contained identical food rewards but had different
costs – each contained a different number of weights (1.5kg/weight). In condition 1, boxes
contained either one, two or three weights. In condition 2, boxes contained zero, one andtwo weights. All boxes needed to be pulled to extract the food. In condition 3, two boxes
with one and two weights, needed to be pulled, while the third box could be accessed
immediately and contained no weight (i.e. “free food”). Results show that monkeys
adjusted their behavior depending on the box’s weight; they prioritized accessing boxes
which had the lowest weights. In conditions 1 and 2, the boxes with the least weight wereaccessed first (38.46% and 37.96% of trials) or second (55.56% and 52.36%),
respectively. In condition 3, they prioritized the “free food” as the first (54.38%) or
second (69.73%) choice. These results suggest that monkeys may not only take energy
expenditure but also other factors such as social conditions into account when ordering
their decisions among different options.

48

P-09

Food transfer occur between closer individuals?
Makiko Take1*, Takakazu Yumoto1
1

Primate Research Institute, Kyoto University, Inuyama, Aichi, Japan* take.makiko.77a@st.kyoto-u.ac.jp

Food transfer is considered as an important opportunity for juveniles to learn dietchoice directly from other group members. Choice of begging target is also importantbecause getting food from others sometimes increases the tension between the individuals.
I hypothesized that juveniles choose the begging target along with the “closeness”. I
investigated with whom juveniles spend more time, and between whom food transfer
events occur in two groups of free-ranging golden-faced saki (Pithecia chrysocephala).
I followed each group and recorded the individual’s behavior, food, and the proximate
individual’s ID every 10 minutes by scan-sampling methods. Every time I observed thefood transfer events, I recorded the individual’s ID and the food. The total observation
time was 159 hours and 54 food transfer events were recorded.
Both in Group A and Group B, old juveniles (OJ) and young juveniles (YJ) tended to
be with individuals of the same age class when they are not eating. When they are eating,
OJ didn’t show any tendency of proximity whereas YJ still tended to be with YJ. Foodtransfer events were initiated only by juveniles, mainly by YJ (94%). In Group A, thebegging from YJ was directed toward the proximate juveniles, whereas in Group B, thebegging from YJ was directed toward an adult male in high frequency.
From those results, it was suggested that the tendency of proximity seemed to be
determined by the juvenile’s age class. On the other hand, it was suggested that thebegging target is not simply chosen by the proximity. In three adult males of Group B,
the most begged male spent the longest time with adult females and spent the shortest
time alone. Such kind of difference of sociality in individual level possibly affect thejuveniles’ choice of begging targets.

49

P-10

Waiting as Infant Care in Wild Chimpanzees
Hiroko Sakuragi1*
1

Wildlife Research Center, Kyoto University, Kyoto, Kyoto, Japan*hrk.sakuragi@gmail.com

In group living mammals, “waiting” or “pausing” in group travel situations has been
described in many species (capuchins: Leca et al., 2003; Meunier, 2008; macaques: Sueur& Petit, 2009; cattle: Ramseyer et al., 2009) including chimpanzees (Pan troglodytes)(Gruber and Zuberbüler, 2013). Although most literatures that refer to such behavior deal
with collective movement, decision-making and coordination, waiting may also be a formof care toward immature individuals that is possibly crucial for their safety and survival.
Yet, waiting has never been investigated in the context of care. Previous studies on
chimpanzee infant care, both maternal and non-maternal, have focused on behaviors thatinvolve body contact (Van-Lawick Goodall, 1968; Plooij, 1984, 1987; Nishida, 1983;Bădescu et al., 2016), which may have led to a biased outlook on caretaking in
chimpanzees. Significance of indirect care has been underestimated across all taxa, whichcalls for a wider approach to the behavioral aspect of infant care (Rosenbaum & Gettler,
2018b). Chimpanzees live in fission-fusion societies, resulting in sub-groups frequently
dissolving when members leave a certain site. Being attended to when travel begins likely
carries great importance to immature individuals in moving within the home range safely.
Waiting for such individuals is, naturally, a task for those who know they are going totravel with them—mothers, and perhaps the youngest siblings of infants. Maternal siblings
are known to be relatively keen on taking care of their younger siblings (Van-Lawick
Goodall, 1968, 1986; Pusey, 1990; Hobaiter et al., 2014; Nakamura & Hosaka, 2015),
although with no quantitative study, to what degree they are reliable as caregivers remains
unknown. The largest focus of this study was to elucidate the little-known pattern ofmaternal and sibling waiting behavior in chimpanzees, particularly to examine its nature
as care. Another was to see if elder offspring (siblings) that continue to travel with theirmothers reduce the mother’s effort in waiting, being “helpers” in infant care, provided
waiting could be considered care. Using GLMM, I examined multiple factors thatpossibly affected the occurrence of maternal and sibling waiting behavior including thoselikely related to the needs of the infant. The results indicated that waiting is indeed care,shown more frequently when infants likely needed it more, primarily by mothers andoccasionally by siblings. It also became apparent that mothers and siblings were likely
aware of each other’s locations and circumstances, ready to adjust their subsequent
behaviors if necessary. Mothers left their infants without waiting when they likely knewthat siblings can attend to them, while siblings likely noticed when mothers have preceded
and waited in their place. Mothers likely utilized their elder offspring to lighten their own
load, while elder offspring responded to situations in which they were needed as
caregivers, acting as infants’ “babysitters” (Kleiman and Malcolm, 1981). As motherswith elder offspring spent less effort in waiting than those without, and among the prior,mothers waited least when said offspring were close by, continued association between
mother and elder offspring could be said to reduce the mothers’ burden in waiting for
infants, elder offspring consequently functioning as “helpers”.

50

P-11

Social grooming in rescued chimpanzees: how early social disruption
and time of rehabilitation affect sociality?
Evelina Daniela Rodrigues1*, and Miquel Llorente2
1

William James Center for Research, ISPA – Instituto Universitário, Lisbon, Portugal
Unitat de Recerca i Etologia, Fundació Mona, Riudellots de la Selva, Girona, Spain.

2*

danielatrodrigues@gmail.com

Rescued chimpanzees living in sanctuaries may have their social skills impaired due totheir traumatic past. Often these chimpanzees were single-housed, and they came frompet or entertainment industry. Grooming is a complex social activity that plays a crucialrole in the maintenance of social relationships. Therefore, the monitorization of thisbehaviour in sanctuaries could be a good indicator of the welfare and sociality vised in arehabilitation process. The aim of this study was to analyse how grooming varies
according to: origin (wild or captive), past circumstance (pet, entertainment or mixed),
previous social housing (single-housed or social-housed) and time of rehabilitation (up toor more than 10 years). For 138.8 hours, we recorded the duration of grooming in 13 adultchimpanzees living at Fundació Mona. Our results indicate that past circumstance andprevious housing condition do not affect grooming duration, but chimpanzees born incaptivity had a higher duration of grooming than chimpanzees born in the wild. Moreover,chimpanzees experiencing more time of rehabilitation have groomed more thanchimpanzees that arrived at Fundació Mona more recently. In conclusion, the higher
duration of grooming seems to be a positive consequence of the success of therehabilitation process. The rehabilitation time may attenuate the differences in grooming
duration that may have been caused by the social disruption factors such as past
circumstance and previous social housing. Nevertheless, the different environments inwhich they have been born seems to still influence the duration of grooming in theindividuals of Fundació Mona.

51

P-12

Reaction of a group of chimpanzees to the death of the oldest female
in the City Zoological Garden of Warsaw
Anna Jakucińska1, Maciej Trojan2*, Julia Sikorska2,3, Katarzyna Wejhert1, Maciej Kapusta1,
Monika Kostrzewa1 and Dominika Farley4
1

2

Warsaw Municipal Zoological Garden, Warsaw, Poland
Department of Ethology and Comparative Psychology, Nicolaus Copernicus University, Toruń, Poland
3
Department of Animal Behaviour, University of Warsaw, Warsaw, Poland
4
Institute of Psychology, Jagiellonian, Kraków, Poland*

julia.sikorska@umk.pl

The data presented here refers to the death of the oldest female in a troop of 8
chimpanzees living in a habitat of the City Zoological Garden of Warsaw (Poland). At
the time of natural death the female (Judy) was 53 years old. The greatest attention paid
to the body, in various ways, was by the youngest member, a 3-year-old male (Frodo).
The females in general spent more time with and touched the body more often than themales did. The females included a daughter of the dead female (Mandy, 23 years old),and a granddaughter (Lucy, 14 years old). One of the males, Szymi (the oldest, at 32 yearsof age), alone avoided all contact with the dead body. Mandy, after the body of her motherwas removed, made her bedding that evening in the spot where the dead female had lain
the previous night.

52

P-13

Spatial representation of age in chimpanzees
Yuri Kawaguchi1*, Ikuma Adachi1, Masaki Tomonaga1,
1

Primate Research Institute, Inuyama, Aichi, Japan*

kawaguchi.yuri.57r@st.kyoto-u.ac.jp

Previous studies have shown that humans associate abstract concepts with space, along
both vertical and horizontal plane. One example is the fact that we have mental timeline
and associate space and time (i.e. earlier is left; later is right). It is also known that certain
spatial representation is shared with non-human primates. For example, one study showedthat high-ranked individuals are represented in higher positions in chimpanzees too. How
about mental timeline, especially individual age in chimpanzees? We investigatedwhether chimpanzees associate space (top-bottom, or left-right) and age (infant-adult)
using matching-to-sample procedure. Six chimpanzees participated the experiment. A
picture of either adult or infant chimpanzee face (“sample”) was appeared on the centerof the monitor upon their touching the start key. The sample disappeared after 750 ms.
Then two choice alternatives were presented on top and bottom (or left and right)
positions. The alternatives were from either same age category or different age categories.
One of them was the same as the sample picture and chimpanzees were required to touchthat. If chimpanzees spontaneously map age category onto space, their response time
would be shorter when the position of correct choice was congruent with their mental
representation. This study is still on-going and we are presenting the result in this
symposium for the first time.

53

P-15

Development of Combinatory Manipulation in Great Apes andHumans: Implication for Action Patterns in Tool Use
Misato Hayashi1*and Hideko Takeshita212Primate Research Institute, Kyoto University, Inuyama, Aichi, Japan
Graduate School of Psychology, Otemon Gakuin University, Ibaraki, Osaka, Japan*hayashi.misato.4e@kyoto-u.ac.jp

Object manipulation can be used as a comparative scale of cognitive development
among primates including humans. Some previous studies focused on object
manipulation and/or tool-using behavior as an indicator of cognitive abilities inchimpanzees and human children. Particularly, combinatory manipulation is a precursorof tool-using behavior and starts at around 10 months of age in humans. Mother-reared
chimpanzees started combinatory manipulation at 8-11 months of age which wascomparable to humans. Inserting action was commonly observed in chimpanzees from anearly age. However, stacking action appeared later in captive chimpanzees’ development.
This pattern may reflect the tool-use repertoire of wild chimpanzees where tool-use
behaviors involving the inserting action are wide spread among most of the wild
chimpanzee populations. Stacking-block behavior started at around one year of age inhumans and at two years and seven months in one out of three chimpanzees. All three
chimpanzees started to stack up blocks at three years and one month following trainings
by humans. After the acquisition of stacking action, both chimpanzees and humanchildren showed similar performances in tasks using blocks of various shapes which were
designed to test their physical understanding. Similarly, both chimpanzees and humanchildren showed trial-and-error strategies in combining cups into a nesting structure.
Direct comparison between humans and captive chimpanzees revealed fundamental
similarities in the domain of physical intelligence. Apart from chimpanzees,
developmental data on object manipulation from other species of great apes remains
limited. Through our preliminary investigation, all species of great apes showed two typesof combinatory manipulation (insertion and stacking) in captive settings with somevariations in the timing and order of development. Among captive great apes, common
ages for the emergence of combinatory manipulation was around 2.5 years. Thus,
compared to this great-apes’ standard age, inserting action in chimpanzees appeared froman exceptionally early age and it was comparable to humans. This early appearance ofcombinatory manipulation in captive chimpanzees may reflect the tool-use commonality
in wild chimpanzees, especially the ones involving the inserting action. Although all four
species of great apes possess the fundamental cognitive ability of performing
combinatory manipulation in captivity, socio/environmental factors may contribute to thetool-use repertoire and frequency in the wild.

54

P-16

An ongoing research:familiar vs. unfamiliar human face recognition in chimpanzees
Yena Kim1*, Jae Chun Choe1, and Masaki Tomonaga2
1

Research Institute of EcoScience, Ewha Womans University, Seoul, Republic of Korea
2
Primate Research Institute, Kyoto University, Inuyama, Japan*

wildlife1984@gmail.com

It is well known that humans extract information about individuals, such as identity,sex, emotional state, etc., from face and use this information for everyday communication.
Previous research on primate face perception as well have revealed similar patterns inprimates, the ability of individual and emotional recognition and configural processing ofconspecific faces. However, studies using an eye-tracker have revealed a major difference
in attentional gaze on face between humans and other nonhuman great apes in whichhumans focus longer time on the eye region contrary to the great apes displaying fastscanning of eyes and mouth regions. These studies suggest that the amount of information
being conveyed from each facial component, such as eyes, a nose and a mouth, can differ
between humans and nonhuman great apes. Although there have been a considerable
amount of research on great ape face perception, no study has yet tested whether thedifferential attentional preference on facial components found in eye-tracking studies canbe further retained to the short- or long-term memories. Furthermore, studies about
whether great apes also process familiar and unfamiliar faces differently are lacking. The
current study was designed to investigate the relative contribution of facial components(eyes, nose, mouth) to familiar vs. unfamiliar human face recognition in chimpanzees,
our closest phylogenetic relative. Eight chimpanzees at PRI have been participating in afull-face to full-face delayed matching-to-sample training with familiar and unfamiliar
face stimuli sets. After the training, 1) full-face to components (eyes, nose, mouth)
matching and 2) memory retrieval (discrimination task) tests will be introduced.

55

P-17

Chimpanzee’s categorical classification of the sound: The voice ofchimpanzee vs pure tone
Hiroya Takiyama1*, Yuko Hattori1 and Masaki Tomonaga1
1Primate Research Institute, Kyoto University, Inuyama, Japan*takiyama.hiroya.23u@st.kyoto-u.ac.jp

Primates use vocalization in their habitat. However, there are little studies of theirauditory cognitive ability. I conducted the experiment with chimpanzees using soundstimuli. In the experiment, chimpanzees distinguish chimpanzees’ voice from pure tone
because to communicate with others using vocalization, it is very important to distinguish
voice of others from non-vocal sounds and understand their meanings and reply to them.
I examined which acoustic features they use when they recognize voice of the otherindividuals. The chimpanzees were required to push the button on the same side of the
speaker which played chimpanzees’ voice, ignoring pure tone played simultaneously
from the other speaker. Although the chimpanzees are under training, accuracy of one offour chimpanzees already reached nearly 90% in fourteen sessions. In the second
experiment, I focused on the effect of vocalization on behavioral response. This
experiment will help to understand chimpanzees’ auditory cognitive ability.

56

P-18

Informing conservation efforts through quantitative assessments ofhabitat suitability for chimpanzees in the Greater Nimba Landscape,Guinea, Africa
Maegan Fitzgerald1*, Robert Coulson2, 3, A. Michelle Lawing2, Tetsuro Matsuzawa4, andKathelijne Koops5
1

Wildlife Research Center, Kyoto University, Kyoto, Japan
Department of Ecosystem Science and Management, Texas A&M University, College
Station, Texas, United States of America
3
Knowledge Engineering Laboratory, Department of Entomology, Texas A&M
University, College Station, Texas, United States of America
4
Kyoto University Institute for Advanced Study, Kyoto, Japan
5
Anthropological Institute and Museum, University of Zurich, Zurich, Switzerland

2*maeg2nimba@gmail.com

In West Africa, the Upper Guinean Forest Ecosystem, is a biodiversity hotspot that has
undergone dramatic deforestation and fragmentation. This ecosystem has been astronghold for many endemic plant and animal species and sustains the livelihoods ofmillions of people across West Africa. While Guinea holds only 6% of the remaining
intact, dense Upper Guinean forest, the country is home to the largest remaining
populations of Western chimpanzees (Pan troglodytes verus), the only sub-species ofchimpanzee listed as Critically Endangered by the IUCN. To effectively address threatsto Western chimpanzees, such as habitat loss/degradation, it is important to understandhow chimpanzees respond to their environment and identify the environmental factors
influencing habitat selection. For this study, species distribution modeling was used to
identify areas of suitable chimpanzee habitat and the biophysical variables contributing
to habitat suitability within the Greater Nimba Landscape (GNL), Guinea. The model
results highlight the importance of dense, healthy vegetation, elevation, topographic
exposure, and proximity to rivers for chimpanzees while also emphasizing that suitability
is isolated and does not make up a large portion of the GNL. Through this research,quantitative, species-specific information can be used to inform broader conservation andland-use planning initiatives within the GNL.

57

P-19

Characterization of bitter taste sensitivity of four species of Sulawesi
Macaques
Xiaochan YAN 1*, Kanthi Arum Widayati 2, 6, Nami Suzuki-Hashido3, Fahri Bajeber4,
Akihiro Itoigawa1, Laurentia Henrieta Permita Sari Purba2, Bambang Suryobroto2,
Yohey Terai5, Hiroo Imai1
1

Primate Research Institute, Kyoto University, Inuyama, Japan;
Department of Biology, Bogor Agricultural University, Bogor, Indonesia;
3
Academy of Emerging Sciences, Chubu University, Kasugai, Japan;
4
Department of Biology, Tadulako University, Palu, Indonesia
5
Department of Evolutionary Studies of Biosystems, The Graduate University for Advanced Studies,
Hayama , Japan
6
Wildlife Research Center, Kyoto University, Kyoto, Japan2*

yanxch3@outlook.com

Sulawesi macaques are divergently evolved into 7 species within a relatively shortperiod and live allopatrically on Sulawesi Island, which is isolated for million years long.
Thus, it will be interesting to study functional genes which might reveal their specificspecies difference. Bitter taste plays an important role in avoiding ingestion of toxins andinduces innate avoidance behaviors, which might evolve to reflect species-specific diets
during mammalian evolution. In addition, it was recently reported that the receptors are
involved in the immuno-system against bacteria and parasites. One of the genes is
TAS2R38, receptor for bitter compound Phenylthiocarbamide (PTC) and some bacterial
secreting compounds. Here, we report characterization of TAS2R38 in four species of
Sulawesi macaque, M. hecki (N: 16, M. tonkeana (N: 12), M. nigrescens (N: 11and M.
nigra (N: 15. We conducted behavioral experiment on PTC acceptance and later
functional analysis on amino acid residue(s) of TAS2R38 responsible for low bitter taste
sensitivity. Our result shows that both M. tonkeana and M. nigra were found with “lowsensitive” but exhibited different pattern in genetic aspect. Amino acid changes at
position 117, 130, 134 of M. tonkeana, whereas one base insertion caused early stopcodon at site 178 of M. nigra, leading to non-taster phenotype separately. By haplotype
network construction, we found that ‘low-sensitive’ haplotypes were independently
evolved after speciation. This finding might give a clue for clarifying evolutionary
relationship and dietary habits among the four species, as bitter taste may be related tothe plants they consume.

58

P-20

Sedative and Antinociceptive Effects of S(+)-Ketamine and Midazolam
or Fentanyl and Droperidol in Black-tufted-ear marmosets
Luiz F. Larangeira1*, Mario A. Ferraro2, Vanessa N. Gris3 and Silvia R.G. Cortopassi2
1

Technical Division of Veterinary Medicine and Wildlife Management,
Municipal Secretariat for Environment of São Paulo, São Paulo, Brazil
2
Department of Veterinary Surgery, University of São Paulo, São Paulo, Brazil
3
Center for Human Evolution Modeling Research, Primate Research Institute, Kyoto University,
Inuyama,Japan*

larangeira.vet@gmail.com

The cardiorespiratory, sedative and antinociceptive effects of the combination of S(+)ketamine with midazolam and fentanyl with droperidol in Callithrix penicillata werecompared in this study. Twenty-six adult black-tufted-ear marmosets, belonging to theTechnical Division of Veterinary Medicine and Wildlife Management of the Secretariat
for Environment of São Paulo, Brazil, underwent chemical immobilization for physicalexamination, blood sampling, tattooing and microchipping. Animals were randomly
treated with S(+)-ketamine (10 mg/kg) and midazolam (1 mg/kg) (KM; BW = 314.6 ±617g) or fentanyl (12,5 µg/kg) e droperidol (625 µg/kg) (FD; BW = 278.9 ± 87.7g), given
by intramuscular injection. Heart rate and rhythm (lead II ECG) and respiratory rate (chest
movement) were recorded. Sedation, antinociception (response to interdigital pinch),muscle relaxation, auditory (a handclap close to the animal’s ears), and visual response
were evaluated using a modified and previously described scoring system. We also
evaluated the sedative onset and effective times, and recovery time and quality. Allvariables were recorded within 5 minutes of administration and at 10-min intervals.
Sedation in KM was achieved faster (KM = 1.95 ± 0.56 min; FD = 6.5 ± 0.66 min; p <0,0001) and lasted for a shorter period of time (KM = 50.7 ± 16.1 min; FD = 79.2 ± 21.8
min; p = 0,0009), although the animals remained in lateral recumbency longer than in FD.
KM was similar to FD in its cardiorespiratory effects, auditory and visual responses but
produced ataxia and excitement during the recovery period (5/13), while FD showed
vocalization (3/13) and one episode of vomiting. FD combination produced less sedationand muscle relaxation but superior degree of nociception compared to KM. The
administration of fentanyl and droperidol resulted in adequate sedation, without clinicallyimportant cardiorespiratory effects, and is suitable for short-term procedures that require
more intense analgesia in black-tufted-ear marmosets.

59

P-21

Inter-group relationships of feral horses (Equus caballus) in Serra
D’Arga, Portugal:
Tamao Maeda1*, Sakiho Ochi1, Monamie Ringhofer2, Satoshi Hirata1 and Shinya Yamamoto2
1
2

Wildlife Research Center, Kyoto University, Kyoto, Japan
Institute for Advanced Study, Kyoto University, Kyoto, Japan*

tamao@powarch.com

Some mammalian species of different taxonomic groups, including humans, havedeveloped social structures with nested levels of organization. This multilevel society
is one of the most complex social systems in animals, but their function and evolutional
process are still poorly understood especially for non-primate species because it requires
good observation of large numbers of identified individuals. Equine groups are one of
the taxa that have nested social structure, and there are some studies on several species
such as zebras and Przewalski's horses, but not on domestic horses (Equus caballus).
Studying the inter-group relationships of feral horses and comparing those of the otherequine species may help understanding their origin and their ecological and social
meanings. In this study, we aimed to reveal whether domestic horses form multilevel
societies. We took aerial photos of feral horse herds in Serra D’Arga Portugal in 30
minutes interval using drones, identified all the individuals and collected their position
data. In the field, we observed 21 harems and 2 bachelor groups and we found somepatterns in the group positioning. This result strongly indicates the multilevel structuresof the feral horse societies.

60

P-23

Do female feral horses (Equus caballus) choose their herd?
Sakiho Ochi1*, Monamie Ringhofer2, Renata Mendonca3 and Satoshi Hirata1
1
2

Wildlife Research Center, Kyoto University, Kyoto, Japan
Institute for Advanced Study, Kyoto University, Kyoto, Japan
3
Primate Research Institute, Kyoto University, Kyoto, Japan*ochisakiho11@gmail.com

The final purpose of this study is to clarify the reason why female feral horses (Equus
caballus) do secondary transfers. In many species, either or both sex of offspring disperses
to other group from their natal group, when they mature, in order to avoid inbreeding(natal disperse). On the other hand, some individuals of those species change their groupagain after natal disperses (secondary inter-group transfer), but it is still poorly
understood. Besides that, it is generally said that female of harem species need not tochoose male. In previous study about feral horse, it is reported that the female is more
attacked by other females of new group when they transfer to other herds. Therefore,changing her group can be cost to female. However, they do not seem to have goodreasons to change their group, we have already observed the secondary transfer 17
examples of 15 individuals of feral horse between 35 herds in Serra D’Arga Portugal
2016-2018. Six examples of that were natal disperses and the other 11 examples weresecondary transfers. In this presentation, I will report about preliminary analysis aboutsome factors which is predicted to related to the secondary transfer and females’ herd
choice, including seasons, size of the group, the number of the males of the herd and therelationship between females.

61

P-24

A study plan about feeding behavior in captive giraffes
Anna Kawakita1*
1

Wildlife Research Center, Kyoto University, Japan*

kawakita.anna.32x@st.kyoto-u.ac.jp

It is said that ruminating behavior allows many kinds of even toed ungulates to digest
food in safe places and spend less time taking it in places where they may be at risk of
predation. Some experimental studies on domestic cattle, goats and sheep show thatrumination occurs more often at night than in the daytime. Most ruminants do it lying
down, while giraffes (giraffa camelopardalis), which are the largest ruminants in theworld, do not lie so often because it takes them longer to stand up if danger arises. Giraffes
chew their cud standing and lying down, but we have little knowledge of the exact detailsof their postures while ruminating. Previous studies show that ruminating behavior ingiraffes could be influenced by food, their feeding behavior, the ambient temperature or
their psychological states. The aim of this study is to investigate feeding behavior ingiraffes including rumination, focusing on their postures, from the observation of captive
giraffes in Japan. In this presentation, I discuss this study plan.

62

P-25

Using dynamic animal social networks to improve the study of
infectious disease transmission
Kenneth Keuk1*, Julie Duboscq1, and Andrew J. J. MacIntosh1
1

Primate Research Institute, Kyoto University, Inuyama, Aichi, Japan*kenneth.keuk@gmail.com

People change, so do their relationships. Social animals also build dynamic
relationships. Social Network Analysis (SNA), a mathematical tool used to summarize
and quantify the relationships in a group at different scales, can be used to study links
between social behaviour and infectious disease transmission. For instance, studying
Japanese macaques on Kōshima (Macaca fuscata) and Yakushima (M. f. yakui) has
shown positive correlations between an individual’s centrality in the social network andits degree of gastro-intestinal parasitism. But since these correlations relied on social
networks derived from aggregate data collected over large periods (e.g. whole seasons),
there was little room for testing whether social relationships change with time, and how
this might affect the relationship between networks and infection. Moreover, current(estimated) levels of parasite infection do not reflect current levels of parasiteacquisition, which is really what we aim to link with social networks. Especially forintestinal helminths, a prepatent period defines the time interval between when parasites
are acquired and when they can be detected in a host, something previous studies havenot considered. The aim of the present study was thus to test whether dynamic social
networks, inclusive of a time lag (7-10 weeks prior to infection measurements) toincorporate the prepatent period of a specific intestinal nematode (Trichuris sp.), better
predict infection phenotypes than static networks derived from aggregate and concurrent
data. We use our model on a simulated extreme example before applying it to an empirical
data set from Kōshima macaques, and discuss the potential implications and applicationsof such models for epidemiological research.

63

The 10th International Symposium on Primatology and Wildlife Science

Workshop on Science Communication:
Mobile Filmmaking
September 23rd (Sun) 2018, 1:00 – 4:00 PM
Science Seminar House, Kyoto Univ. (North Campus)

Dr. Wiebke Finkler

Prof. Lloyd Spencer Davis

Department of Tourism,
University of Otago

Stuart Professor of Science Communication,
University of Otago

Would you like to learn how to create engaging science communication content?
This is a practical workshop for scientists, researchers and postgraduate students wanting toconduct science communication/ educational outreach with a focus on how to create
engaging multimedia content for public communication and sharing online. It has a particularfocus on video production and storytelling. The workshop will introduce marketing andscience communication theory and practice as they apply to effective communication ofscientific research. Core facets of the workshop include:• Elements required for effective science communication and marketing communication• Storytelling• Creating engaging and contagious communication and educational outreach• Packaging science content• Social media• Using filmmaking as outreach and/or research tools
NOTE: bring your own tablet or smartphone capable of taking videos for some hands-on work.

✉pws_sympo2018_admin@pws.wrc.kyoto-u.ac.jp

Conserv’Session
Conservation Documentary Series

保護についてのドキュメンタリーシリーズ

Special Guest:
スペシャルゲスト

Prof. Fred Bercovitch
Specially-appointed Professor of
Kyoto University, giraﬀe expert
and Execu<ve Director of Save theGiraﬀeswww.savethegiraﬀes.org

2018年9月24日（月）September 24th (Mon) 2:00-5:30 PM
どなたでも無料で参加いただけます!
No registraHon required, free entry!
音声：英語字幕（英語字幕）
Language: English (English subHtles)
Discussion: English/Japaneseconservsession@googlegroups.com
www.facebook.com/ConservSession
twitter.com/ConservSession

京都大学北部キャンパス 理学部セミナーハウス
Science Seminar House
Kyoto University (North Campus)
Schedule
2:00-3:00 Giraffes: Africa's Gentle Giants
3:00-3:15 Break
3:15-3:45 Last of the Longnecks
3:45-5:30 Discussion

/ŶƚƌŽĚƵĐƚŝŽŶƚŽƚŚĞWƌŽŐƌĂŵ
>ĞĂĚŝŶŐ'ƌĂĚƵĂƚĞWƌŽŐƌĂŵŝŶWƌŝŵĂƚŽůŽŐǇĂŶĚtŝůĚůŝĨĞ^ĐŝĞŶĐĞ;Wt^ͿdŚĞ >ĞĂĚŝŶŐ 'ƌĂĚƵĂƚĞ WƌŽŐƌĂŵ ŝŶ WƌŝŵĂƚŽůŽŐǇ ĂŶĚ tŝůĚůŝĨĞ ^ĐŝĞŶĐĞ ;Wt^Ϳ ƐƚƌŝǀĞƐ ƚŽ ƌĞĂůŝǌĞ ŐůŽďĂů ǁĞůůͲďĞŝŶŐ͘ dŚĞ ƉƌŽŐƌĂŵ ĂŝŵƐ ƚŽ ĨŽƐƚĞƌ ŝŶĚŝǀŝĚƵĂůƐ
ǁŝƚŚ ƚŚĞ ĂďŝůŝƚǇ ƚŽ ŵĂŬĞ ƋƵŝĐŬ ũƵĚŐĞŵĞŶƚƐ ĂďŽƵƚ ĞŶǀŝƌŽŶŵĞŶƚĂů ŝƐƐƵĞƐ ĂŶĚ ƚŽ ĚĞƐŝŐŶ Ă ĨƵƚƵƌĞ ŐůŽďĂů ƐŽĐŝĞƚǇ͕ ǁŚŝůĞ Ăƚ ƚŚĞ ƐĂŵĞ ƚŝŵĞ ŶƵƌƚƵƌŝŶŐ ůĞĂĚĞƌƐ
ŝŶĚŝƐƉĞŶƐĂďůĞ ĨŽƌ ŐůŽďĂů ĂĐƚŝŽŶ͘:ĂƉĂŶĞƐĞ ƉƌŝŵĂƚŽůŽŐǇ ŚĂƐ ƉůĂǇĞĚ Ă ůĞĂĚŝŶŐ ƌŽůĞ ŝŶ ƚŚŝƐ ƵŶŝƋƵĞ ĂĐĂĚĞŵŝĐ ĞŶĚĞĂǀŽƌ͘ ƵƌŝŶŐ ƌĞĐĞŶƚ ǇĞĂƌƐ͕ ƚŚĞ ĨŝĞůĚ ŽĨ ͞tŝůĚůŝĨĞ ^ĐŝĞŶĐĞ͕͟ ǁŚŝĐŚ ƚĂƌŐĞƚƐ
ĞŶĚĂŶŐĞƌĞĚ ƐƉĞĐŝĞƐ ƌĞƐĞĂƌĐŚ͕ ŚĂƐ ďĞŐƵŶ ƚŽ ĞŵĞƌŐĞ͘ tŝƚŚ ĨŝĞůĚǁŽƌŬ ĂƐ ŝƚƐ ĨŽƵŶĚĂƚŝŽŶ͕ ƚŚĞ ĨƵŶĚĂŵĞŶƚĂů ĂŝŵƐ ŽĨ ƚŚŝƐ ĨŝĞůĚ ĂƌĞ Ă ĐŽŵƉƌĞŚĞŶƐŝǀĞ
ƵŶĚĞƌƐƚĂŶĚŝŶŐ ŽĨ ƚŚĞ ŚƵŵĂŶ ŵŝŶĚ͕ ďŽĚǇ͕ ůŝĨĞ ĂŶĚ ŐĞŶŽŵĞ͕ ĂƐ ǁĞůů ĂƐ ŚĂŶĚƐͲŽŶ ĂĐƚŝǀŝƚŝĞƐ ƚŚĂƚ ƚĂƌŐĞƚ ƚŚĞ ǁĞůůͲďĞŝŶŐ ŽĨ ƚŚĞ ǁŽƌůĚ͘tŚŝůĞ ŽŶ ƚŚĞ ĨƌŽŶƚůŝŶĞƐ ŽĨ ĂĐĂĚĞŵŝĐ ƌĞƐĞĂƌĐŚ͕ ǁŝůĚůŝĨĞ ƐĐŝĞŶĐĞ ŝŶ :ĂƉĂŶ ůĂĐŬƐ ƚŚƌĞĞ ŝŵƉŽƌƚĂŶƚ ĐĂƌĞĞƌƐ ƚŚĂƚ ĂůƌĞĂĚǇ ĞǆŝƐƚ ŝŶ ƚŚĞ tĞƐƚ͗ ;ϭͿ ŽŶƐĞƌǀĂƚŝŽŶ
ƐƉĞĐŝĂůŝƐƚƐ ǁŝƚŚ ŝŶƚĞƌŶĂƚŝŽŶĂů ŽƌŐĂŶŝǌĂƚŝŽŶ͕ ƐƵĐŚ ĂƐ ƚŚĞ hŶŝƚĞĚ EĂƚŝŽŶƐ ĂŶĚ ŽƚŚĞƌ E'KƐ͖ ;ϮͿ ƵƌĂƚŽƌƐ ŽĨ ŵƵƐĞƵŵƐ͕ ǌŽŽƐ͕ ĂƋƵĂƌŝƵŵƐ͕ ĂŶĚ ƐŝŵŝůĂƌ
ŝŶƐƚŝƚƵƚŝŽŶƐ͕ ƉĂƌƚŝĐƵůĂƌůǇ ƚŚŽƐĞ ƚŚĂƚ ĐĂŶ ĚĞǀĞůŽƉ ĂŶĚͬŽƌ ĞǆƉĂŶĚ ŵƵƐĞƵŵƐ Žƌ ǌŽŽƐ ĂƐ Ă ͞ĨŝĞůĚ ŵƵƐĞƵŵƐ͟ ŝŶ Ă ƐƉĞĐŝĨŝĐ ŚĂďŝƚĂƚ͖ ĂŶĚ ;ϯͿ ĞĚŝĐĂƚĞĚ
ŝŶĚŝǀŝĚƵĂůƐ ƚŚĂƚ ŝŶǀĞƐƚ ŐƌĞĂƚ ůĞŶŐƚŚƐ ŽĨ ƚŝŵĞ ŝŶ ŽƵƚƌĞĂĐŚ ĂĐƚŝǀŝƚŝĞƐ ŝŶ ƐƉĞĐŝĨŝĐ ĐŽƵŶƚƌŝĞƐ ĂŶĚ ƐŽĐŝĞƚŝĞƐ͕ ƚŚĞ ƐŽͲĐĂůůĞĚ ͞ďŽŽƚƐͲŽŶͲƚŚĞͲŐƌŽƵŶĚ͟ ĂƉƉƌŽĂĐŚ͘tŚŝůĞ ƉƌŽǀŝĚŝŶŐ Ă ĨŽƵŶĚĂƚŝŽŶ ĨŽƌ ŶĞǁ ƌĞƐĞĂƌĐŚ͕ ĞĚƵĐĂƚŝŽŶ ĂŶĚ ŚĂŶĚƐͲŽŶ ĞǆƉĞƌŝĞŶĐĞ͕ ƚŚŝƐ ƉƌŽŐƌĂŵ ĂŝŵƐ ƚŽ ŶƵƌƚƵƌĞ ŐůŽďĂů ůĞĂĚĞƌƐ ǁŚŽƐĞ
ĂĐĐŽŵƉůŝƐŚŵĞŶƚƐ ŐƌŽǁ ŚĂŶĚͲŝŶͲŚĂŶĚ ǁŝƚŚ ƚŚĞ ůĂƌŐĞƌ ĂĐĂĚĞŵŝĐ ĨŝĞůĚ͘ &Žƌ ĨƵƌƚŚĞƌ ĚĞƚĂŝůƐ ŽŶ ƚŚĞ ƉƌŽŐƌĂŵ͕ ƉůĞĂƐĞ ƌĞĨĞƌ ƚŽ ŽƵƌ ,W͘䠄ŚƚƚƉ͗ͬͬǁǁǁ͘ǁŝůĚůŝĨĞͲƐĐŝĞŶĐĞ͘ŽƌŐͬ䠅

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WƌŝŵĂƚĞZĞƐĞĂƌĐŚ/ŶƐƚŝƚƵƚĞ

㼀㼔㼑㻌㼑㼚㼞㼛㼘㼘㼙㼑㼚㼠㻌㼜㼞㼛㼏㼑㼟㼟㻌㼛㼒㻌㼠㼔㼑㻌㻸㼑㼍㼐㼕㼚㼓㻌㻳㼞㼍㼐㼡㼍㼠㼑㻌㻼㼞㼛㼓㼞㼍㼙㻌㼕㼚㻌㻼㼞㼕㼙㼍㼠㼛㼘㼛㼓㼥㻌㼍㼚㼐㻌㼃㼕㼘㼐㼘㼕㼒㼑㻌㻿㼏㼕㼑㼚㼏㼑dŚĞ >ĞĂĚŝŶŐ 'ƌĂĚƵĂƚĞ WƌŽŐƌĂŵ ŝŶ WƌŝŵĂƚŽůŽŐǇ ĂŶĚ tŝůĚůŝĨĞ ^ĐŝĞŶĐĞ ;Wt^Ϳ ŝƐ Ă ϱͲǇĞĂƌ ƉƌŽŐƌĂŵ͘ ^ƚƵĚĞŶƚƐ ĂƉƉƌŽǀĞĚ ƚŽ ũŽŝŶ ƚŚĞ Wt^ ƉƌŽŐƌĂŵ ĨƌŽŵ ƚŚĞŝƌ
ĨŝƌƐƚ ǇĞĂƌ ŽĨ DĂƐƚĞƌ͛Ɛ ƉƌŽŐƌĂŵ ǁŝůů ƉƌŽŐƌĞƐƐ ĨƌŽŵ >ϭ͕ >Ϯ͕ >ϯ͕ >ϰ͕ >ϱ͕ ĂŶĚ ǁŝůů ĐŽŵƉůĞƚĞ ƚŚĞ ƉƌŽŐƌĂŵ ŝŶ ϱ ǇĞĂƌƐ͘dŚĞ Wt^ ƉƌŽŐƌĂŵ ŝƐ ĐŽŵƉůĞƚĞĚ ďǇ ƐƚƵĚĞŶƚƐ ƉĂƌĂůůĞů ƚŽ ƚŚĞŝƌ ĞǆŝƐƚŝŶŐ <ǇŽƚŽ hŶŝǀĞƌƐŝƚǇ ŵĂƐƚĞƌ͛Ɛ ĂŶĚ ĚŽĐƚŽƌĂů ƉƌŽŐƌĂŵƐ͘ dŚĞƌĞĨŽƌĞ͕ ƐƚƵĚĞŶƚƐ ĚŽ ŶŽƚŶĞĞĚ ƚŽ ĐŚĂŶŐĞ ƚŚĞŝƌ ƐƵƉĞƌǀŝƐŽƌ Žƌ ƐĞĐƚŝŽŶͬůĂďŽƌĂƚŽƌǇ ƚŽ ũŽŝŶ Wt^͘ ,ŽǁĞǀĞƌ͕ ƚŚĞƌĞ ĂƌĞ ƚǁŽ ŶĞĐĞƐƐĂƌǇ ĐŽŶĚŝƚŝŽŶƐ ĨŽƌ ĞůŝŐŝďŝůŝƚǇ͗ϭ͘  ŐƌĂĚƵĂƚĞ ƐƚƵĚĞŶƚ ŽĨ <ǇŽƚŽ hŶŝǀĞƌƐŝƚǇ͗/ƚ ŝƐ ƌĞƋƵŝƌĞĚ ƚŽ ďĞĐŽŵĞ Ă ŐƌĂĚƵĂƚĞ ƐƚƵĚĞŶƚ ŽĨ ƚŚĞ ŝǀŝƐŝŽŶ ŽĨ ŝŽůŽŐŝĐĂů ^ĐŝĞŶĐĞ͕ 'ƌĂĚƵĂƚĞ ^ĐŚŽŽů ŽĨ ^ĐŝĞŶĐĞ ;<ǇŽƚŽ hŶŝǀĞƌƐŝƚǇͿ͘ ,ŽǁĞǀĞƌ͕ ǁĞ ĂƌĞ ŝŶ ƚŚĞƉƌŽĐĞƐƐ ŽĨ ĂĚũƵƐƚŵĞŶƚ ĨŽƌ ƐƚƵĚĞŶƚƐ ŽĨ ŽƚŚĞƌ ŐƌĂĚƵĂƚĞ ĚĞƉĂƌƚŵĞŶƚƐ ƚŽ ĞŶƌŽůů ŝŶ ŽƵƌ ƉƌŽŐƌĂŵ͕ ƐŽ ƉůĞĂƐĞ ĚŽ ŶŽƚ ŚĞƐŝƚĂƚĞ ƚŽ ŝŶƋƵŝƌĞ͘Ϯ͘ dŽ ĂƉƉůǇ ĂŶĚ ƌĞĐĞŝǀĞ ĂƉƉƌŽǀĂů ƚŽ ĞŶƌŽůů ŝŶƚŽ ŽƵƌ ƉƌŽŐƌĂŵ͗dŚĞ ƉƌŽĐĞƐƐ ŝƐ ƚŚĞ ƐĂŵĞ ĨŽƌ ďŽƚŚ :ĂƉĂŶĞƐĞ ŶĂƚŝǀĞƐ ĂŶĚ ĨŽƌĞŝŐŶ ƐƚƵĚĞŶƚƐ͘ ůŝŐŝďůĞ ƐƚƵĚĞŶƚƐ͗ ϭƐƚ ǇĞĂƌ DĂƐƚĞƌ͛Ɛ ƐƚƵĚĞŶƚƐ ;ǁŝůů ďĞ ĐĂůůĞĚ >ϭ ƐƚƵĚĞŶƚͿ͕ Žƌ ĂĚŽĐƚŽƌĂů ƐƚƵĚĞŶƚƐ ;ǁŝůů ďĞ ĐĂůůĞĚ >ϯ ƐƚƵĚĞŶƚͿ͘ ŶŶƵĂůůǇ͕ ǁĞ ǁŝůů ĚŝƐĐůŽƐĞ ƚŚĞ ŐƵŝĚĞůŝŶĞƐ ĨŽƌ ĂƉƉůŝĐĂŶƚƐ ŝŶ ŵŝĚͲ:ĂŶƵĂƌǇ͕ ĂŶĚ ĂĚŵŝŶŝƐƚĞƌ ƚŚĞ ĞŶƚƌĂŶĐĞ ĞǆĂŵŝŶ ƚŚĞ ďĞŐŝŶŶŝŶŐ ŽĨ DĂƌĐŚ͘ dŚĞ ƐĐŚĞĚƵůĞ ĨŽƌ KĐƚŽďĞƌͲĞŶƌŽůůĞĚ ƐƚƵĚĞŶƚƐ ŝƐ ĚŝĨĨĞƌĞŶƚ͘ ;ŚƚƚƉ͗ͬͬǁǁǁ͘ǁŝůĚůŝĨĞͲƐĐŝĞŶĐĞ͘ŽƌŐͬŝŶĚĞǆͲĞŶ͘ŚƚŵůͿƵƌƌŝĐƵůƵŵ
>ĞĂĚŝŶŐ'ƌĂĚƵĂƚĞWƌŽŐƌĂŵŝŶWƌŝŵĂƚŽůŽŐǇĂŶĚtŝůĚůŝĨĞ^ĐŝĞŶĐĞ;Wt^ͿdŚĞĨŽůůŽǁŝŶŐĐŽŶƚĞŶƚƐƐŚŽǁƚŚĞĐƵƌƌŝĐƵůƵŵĂŶĚƐĐŚĞĚƵůĞƚŚĂƚƚŚĞ>ĞĂĚŝŶŐ'ƌĂĚƵĂƚĞWƌŽŐƌĂŵŝŶWƌŝŵĂƚŽůŽŐǇĂŶĚtŝůĚůŝĨĞ^ĐŝĞŶĐĞ ŽĨĨĞƌ͘ ƌĞĚŝƚƐ
ŽďƚĂŝŶĞĚ ƚŚƌŽƵŐŚ ƚŚĞŵĂŶĚĂƚŽƌǇĐŽƵƌƐĞƐĐĂŶĂůƐŽďĞƵƐĞĚĂƐĐƌĞĚŝƚĨŽƌ'ƌĂĚƵĂƚĞ^ĐŚŽŽůŽĨ^ĐŝĞŶĐĞ͕<ǇŽƚŽhŶŝǀĞƌƐŝƚǇ͘
EKd͗ŽƵƌƐĞƐĐŚĞĚƵůĞƐĂƌĞƐƵďũĞĐƚƚŽĐŚĂŶŐĞ͘&ŽƌŵŽƌĞŝŶĨŽƌŵĂƚŝŽŶ͕ƌĞĨĞƌƚŽƚŚĞĨŽůůŽǁŝŶŐƐŝƚĞ͗ŚƚƚƉ͗ͬͬǁǁǁ͘ǁŝůĚůŝĨĞͲƐĐŝĞŶĐĞ͘ŽƌŐͬĞŶͬĐƵƌƌŝĐƵůƵŵͬWůĞĂƐĞĐŽŶƚĂĐƚƚŚĞĨŽůůŽǁŝŶŐĞͲŵĂŝůĂĚĚƌĞƐƐĨŽƌĂŶǇƋƵĞƐƚŝŽŶƐĂďŽƵƚƚŚĞĐƵƌƌŝĐƵůƵŵŽĨWt^͗ŝŶĨŽΛǁŝůĚůŝĨĞͲƐĐŝĞŶĐĞ͘ŽƌŐ

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EĂƚŝŽŶĂů WĂƌŬ͕ hŐĂŶĚĂ
&Ğď͘ ϭƐƚʹ :Ƶů͘ ϮϭƐƚ͕ ϮϬϭϳ ;>ϰΛZKE'KͿ͗ ƐĞǆ ĚŝĨĨĞƌĞŶĐĞ ŽĨ ŵŽƚŚĞƌ ŽĨĨƐƉƌŝŶŐ ƌĞůĂƚŝŽŶƐŚŝƉƐ ŝŶďŽŶŽďŽ ƉĂƚƌŝůŝŶĞĂů ƐŽĐŝĞƚŝĞƐ
&Ğď͘ ϭϲƚŚʹ :ƵŶĞ ϮϭƐƚ͕ ϮϬϭϳ ;>ϰΛ/E/Ϳ͗sŽĐĂů ĐŽŵŵƵŶŝĐĂƚŝŽŶ ŝŶ ƐŝĂŶ ĞůĞƉŚĂŶƚƐDĂǇ ϴƚŚʹ ^ĞƉ͘ ϭϭƚŚ͕ ϮϬϭϳ ;>ϱΛdĂŶǌĂŶŝĂͿ͗^ŽĐŝĂů ƐƚƌƵĐƚƵƌĞ ĂŶĚ ďĞŚĂǀŝŽƌ ŽĨ ŐŝƌĂĨĨĞ ;ŐŝƌĂĨĨĞ
ĐĂŵĞůŽƉĂƌĚĂůŝƐͿ ŝŶ <ĂƚĂǀŝ EĂƚŝŽŶĂů WĂƌŬ㼀㼛㻌㼐㼑㼢㼑㼘㼛㼜㻌㼟㼗㼕㼘㼘㼟㻌㼕㼚㻌㼜㼘㼍㼚㼚㼕㼚㼓㻌㼜㼞㼛㼖㼑㼏㼠㼟㻌㼀㼛㻌㼘㼑㼍㼞㼚㻌㼟㼡㼞㼢㼕㼢㼍㼘㻌㼟㼗㼕㼘㼘㼟㻌㼍㼟㻌㼠㼔㼑㻌㼎㼍㼟㼕㼟㻌㼍㼕㼙㼑㼐㻌㼍㼠㻌㼛㼚㼑㻌㼛㼞㻌㼙㼛㼞㼑㻌㼛㼒㻌㼠㼔㼑㻌㼠㼔㼞㼑㼑㻌㼑㼤㼕㼠㻌㼒㼛㼞㻌㼒㼡㼠㼡㼞㼑㻌㼒㼕㼑㼘㼐㼣㼛㼞㼗㻚㻌㻭㼏㼠㼕㼢㼕㼠㼕㼑㼟㻌㼕㼚㼏㼘㼡㼐㼑㻦㼜㼛㼕㼚㼠㼟㻌㻔㼓㼛㼍㼘㼟㻕㻌㼛㼒㻌㼠㼔㼑㻌㻼㼃㻿㻌㼜㼞㼛㼓㼞㼍㼙㻌㻔㼕㻚㼑㻚㻘㻌㻙 㼃㼕㼘㼐㼘㼕㼒㼑㻌㼛㼎㼟㼑㼞㼢㼍㼠㼕㼛㼚㼏㼛㼚㼟㼑㼞㼢㼍㼠㼕㼛㼚㻌㼟㼜㼑㼏㼕㼍㼘㼕㼦㼍㼠㼕㼛㼚㻘㻌㼏㼡㼞㼍㼠㼕㼛㼚㻘㻌㻙 㻯㼘㼕㼙㼎㼕㼚㼓㻌㻴㼕㼡㼏㼔㼕 㻹㼛㼡㼚㼠㼍㼕㼚㻌㻔㻞㻘㻠㻞㻜㼙㻕㼛㼡㼠㼞㼑㼍㼏㼔㻕㻚㻌㻾㼑㼝㼡㼕㼞㼑㼐㻌㼠㼛㻌㼐㼑㼟㼕㼓㼚㻛㼏㼛㼚㼐㼡㼏㼠㻌㻙 㻺㼕㼓㼔㼠㻙㼠㼕㼙㼑㻌㼎㼕㼢㼛㼡㼍㼏㻌㼜㼞㼍㼏㼠㼕㼏㼡㼙㻌㼕㼚㼐㼕㼢㼕㼐㼡㼍㼘㻌㼛㼢㼑㼞㼟㼑㼍㼟㻌㼠㼞㼍㼕㼚㼕㼚㼓㻌㼜㼞㼛㼖㼑㼏㼠㼟㻚㻌㻔㼕㼙㼜㼞㼛㼢㼕㼟㼑㼐㻌㼑㼚㼏㼍㼙㼜㼙㼑㼚㼠㻕㻼㼘㼍㼏㼑㻦㻌㻷㼥㼛㼠㼛㻌㼁㼚㼕㼢㼑㼞㼟㼕㼠㼥㻌㻿㼍㼟㼍㼓㼍㼙㼕㼚㼑㻴㾇㼠㼠㼑 㻔㼏㼍㼎㼕㼚㻕㻌㼕㼚㻌㻹㼥㼛㼗㼛㻙㼗㼛㼓㼑㼚㻔㼜㼘㼍㼠㼑㼍㼡㻌㼍㼠㻌㻝㻘㻟㻜㻜㼙㻌㼑㼘㼑㼢㼍㼠㼕㼛㼚㻕㻘㻌㻺㼕㼓㼍㼠㼍DĂŶǇ ŐƌŽƵƉ ĨŝĞůĚǁŽƌŬƐ ĂƌĞ ĚĞƐŝŐŶĞĚ ďǇ ^ƚƵĚĞŶƚƐ͘㻼㼞㼑㼒㼑㼏㼠㼡㼞㼑Ͳ :/'K<hE/ͬd<^</zD㻺㼛㼚㻙㼟㼚㼛㼣㻌㻿㼑㼍㼟㼛㼚㻦㻌Ͳ <ŝĚƐ :ĂŵďŽƌĞĞΛ dK<zK㻶㼡㼘㻚㻌㻝㻣㼠㼔㻙㻞㻜㼠㼔㻘㻌㻞㻜㻝㻤Ͳ ^,KK^,/D㻿㼚㼛㼣㻌㻿㼑㼍㼟㼛㼚㻦㻌Ͳ ^,/ZdK<K ;<ŝůůĞƌ tŚĂůĞͿ㻹㼍㼞㻚㻌㻝㻢㼠㼔㻙㻞㻜㼠㼔㻘㻌㻞㻜㻝㻥Ͳ ^ĂŶ &ƌĂŶĐŝƐĐŽ䕰 㻸㻻㻺㻳㻙
㻸㻻㻺㻳㻙㼀㻱㻾㻹㻌㻵㻺㼀㻱㻾㻺㻿㻴㻵㻼㻌㼀㻾㻭㻵㻺㻵㻺㻳㻿㻌㻔㼏㼛㼞㼞㼑㼟㼜㼛㼚㼐㼟㻌㼠㼛㻌㻰㼛㼏㼠㼛㼞㼍㼘㻌㼏㼛㼡㼞㼟㼑㻕㻔㼏㼛㼞㼞㼑㼟㼜㼛㼚㼐㼟㻌㼠㼛㻌㻰㼛㼏㼠㼛㼞㼍㼘㻌㼏㼛㼡㼞㼟㼑㻕䕰 㻸㻵㻮㻱㻾㻭㻸㻌㻭㻾㼀㻿㻌㻿㼁㻮㻶㻱㻯㼀㻿䖂㻯㼛㼚㼟㼑㼞㼢㼍㼠㼕㼛㼚㻌㻮㼕㼛㼘㼛㼓㼥㻯㼛㼚㼟㼑㼞㼢㼍㼠㼕㼛㼚㻌㻮㼕㼛㼘㼛㼓㼥㻵㼚㼠㼑㼞㼚㼟㼔㼕㼜㻌㼀㼞㼍㼕㼚㼕㼚㼓䖂㻭㼚㼕㼙㼍㼘㻌㼃㼑㼘㼒㼍㼞㼑㻌㻌㻵㼚㼠㼑㼞㼚㼟㼔㼕㼜㻌㼀㼞㼍㼕㼚㼕㼚㼓䖂㻿㼛㼏㼕㼍㼘㻌㻻㼡㼠㼞㼑㼍㼏㼔㻌㻵㼚㼠㼑㼞㼚㼟㼔㼕㼜㻌㼀㼞㼍㼕㼚㼕㼚㼓䖂㻮㼡㼐㼐㼔㼍㻌㻿㼑㼙㼕㼚㼍㼞㼁㻺㻙㼞㼑㼘㼍㼠㼑㼐㻌㼛㼞㼓㼍㼚㼕㼦㼍㼠㼕㼛㼚㼟㻌㼍㼚㼐㻌㻺㻳㻻㼟㻹㼡㼟㼑㼡㼙㼟㻘㻌㼆㼛㼛㼟㻌㼍㼚㼐㻌㻭㼝㼡㼍㼞㼕㼡㼙㼟㻻㼡㼠㼞㼑㼍㼏㼔㻌㼍㼏㼠㼕㼢㼕㼠㼕㼑㼟㻌㼕㼚㻌㼟㼜㼑㼏㼕㼒㼕㼏㻌㼏㼛㼡㼚㼠㼞㼕㼑㼟㻌㼍㼚㼐㻌㼟㼛㼏㼕㼑㼠㼕㼑㼟䕰 㻸㻭㻺㻳㼁㻭㻳㻱㻌㻸㻱㻭㻾㻺㻵㻺㻳䇾㻿㼑㼘㼒㻙
䇾㻿㼑㼘㼒㻙㻿㼠㼡㼐㼥㻌㻼㼍㼞㼍㼐㼕㼓㼙䇿䇾㻴㼍㼚㼐㼟㻙
䇾㻴㼍㼚㼐㼟㻙㼛㼚㻌㻱㼤㼜㼑㼞㼕㼑㼚㼏㼑㻌㼠㼔㼞㼛㼡㼓㼔㻌㻲㼕㼑㼘㼐㼣㼛㼞㼗䇿㻿㼠㼡㼐㼑㼚㼠㼟㻌㼍㼞㼑㻌㼞㼑㼝㼡㼕㼞㼑㼐㻌㼠㼛㻌㼎㼑㼏㼛㼙㼑㻌㼜㼞㼛㼒㼕㼏㼕㼑㼚㼠㻌㼕㼚㻌㼍㼠㻌㼘㼑㼍㼟㼠㻌㼛㼚㼑㻌㼒㼛㼞㼑㼕㼓㼚㻌㼘㼍㼚㼓㼡㼍㼓㼑㻌㼕㼚㻌㼍㼐㼐㼕㼠㼕㼛㼚㻌㼠㼛㻌㼠㼔㼑㼕㼞㻌㼚㼍㼠㼕㼢㼑㻌㼘㼍㼚㼓㼡㼍㼓㼑㻚㻌㻱㼚㼓㼘㼕㼟㼔㻌㼕㼟㻌㼞㼑㼝㼡㼕㼞㼑㼐㻌㼒㼛㼞㻌㼍㼘㼘㻌㼟㼠㼡㼐㼑㼚㼠㼟㻌㼣㼔㼛㼟㼑㻌㼚㼍㼠㼕㼢㼑㻌㼘㼍㼚㼓㼡㼍㼓㼑㻌㼕㼟㻌㼚㼛㼠㻌㻱㼚㼓㼘㼕㼟㼔㻚㻌㻵㼚㼠㼑㼞㼚㼍㼠㼕㼛㼚㼍㼘㻌㼟㼠㼡㼐㼑㼚㼠㼟㻌㼣㼔㼛㼟㼑㻌㼚㼍㼠㼕㼢㼑㻌㼘㼍㼚㼓㼡㼍㼓㼑㻌㼕㼟㻌㻱㼚㼓㼘㼕㼟㼔㻌㼍㼞㼑㻌㼞㼑㼝㼡㼕㼞㼑㼐㻌㼠㼛㻌㼙㼍㼟㼠㼑㼞㻌㼍㼚㼛㼠㼔㼑㼞㻌㼘㼍㼚㼓㼡㼍㼓㼑㻌㼛㼒㻌㼠㼔㼑㼕㼞㻌㼏㼔㼛㼕㼏㼑㻚㻌㻿㼠㼡㼐㼑㼚㼠㼟㻌㼍㼞㼑㻌㼍㼘㼟㼛㻌㼟㼠㼞㼛㼚㼓㼘㼥㻌㼞㼑㼏㼛㼙㼙㼑㼚㼐㼑㼐㻌㼠㼛㻌㼘㼑㼍㼞㼚㻌㼍㻌㼟㼑㼏㼛㼚㼐㻌㼒㼛㼞㼑㼕㼓㼚㻌㼘㼍㼚㼓㼡㼍㼓㼑㻚㻌㻙 㻸㼑㼏㼠㼡㼞㼑㼟㻌㼒㼞㼛㼙㻌㼃㼃㻲㻌㼛㼒㼒㼕㼏㼑㼞㼟㻘㻌㼍㼙㼎㼍㼟㼟㼍㼐㼛㼞㼟㻘㻌㼓㼛㼢㼑㼞㼚㼛㼞㼟㻘㻌㼑㼠㼏㻚㻙 㻻㼒㼒㼕㼏㼕㼍㼘㻌㼘㼍㼚㼓㼡㼍㼓㼑㻦㻌㼚㼛㼠㻌㼟㼜㼑㼏㼕㼒㼕㼑㼐䖂㻭㼟㼡㼞㼍 㻵㼚㼠㼑㼞㼚㼍㼠㼕㼛㼚㼍㼘㻿㼑㼙㼕㼚㼍㼞㻙 㻸㼑㼏㼠㼡㼞㼑㼟㻌㼒㼞㼛㼙㻌㼞㼑㼟㼑㼍㼞㼏㼔㼑㼞㼟㻘㻌㼓㼛㼢㼑㼞㼚㼙㼑㼚㼠㻌㼛㼒㼒㼕㼏㼕㼍㼘㼟㻌㼒㼞㼛㼙㻌㼠㼔㼑㻌㼁㼚㼕㼠㼑㼐㻌㻷㼕㼚㼓㼐㼛㼙㻘㻌㻯㼛㼚㼓㼛㻘㻌㻮㼞㼍㼦㼕㼘㻘㻌㻮㼔㼡㼠㼍㼚㻘㻌㼑㼠㼏㻚㻙 㻻㼒㼒㼕㼏㼕㼍㼘㻌㼘㼍㼚㼓㼡㼍㼓㼑㻦㻌㻱㼚㼓㼘㼕㼟㼔㼀㼔㼑㻌㻵㼚㼠㼑㼞㼚㼍㼠㼕㼛㼚㼍㼘㻌㻿㼥㼙㼜㼛㼟㼕㼡㼙㼟㻌䕰 㼀㼔㼑㻌㻵㼚㼠㼑㼞㼚㼍㼠㼕㼛㼚㼍㼘㻌㻿㼥㼙㼜㼛㼟㼕㼡㼙㼟㻌㼛㼚㻌㻼㼞㼕㼙㼍㼠㼛㼘㼛㼓㼥㻌㼍㼚㼐㻌㼃㼕㼘㼐㼘㼕㼒㼑㻌㼛㼚㻌㻼㼞㼕㼙㼍㼠㼛㼘㼛㼓㼥㻌㼍㼚㼐㻌㼃㼕㼘㼐㼘㼕㼒㼑㻌㻿㼏㼕㼑㼚㼏㼑㻼㼞㼕㼙㼍㼠㼛㼘㼛㼓㼥㻌㼍㼚㼐㻌㼃㼕㼘㼐㼘㼕㼒㼑㻌㻿㼏㼕㼑㼚㼏㼑
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㼀㼔㼑 㻝㼟㼠㻦 㻹㼍㼞㻚 㻜㻢㻙㻜㻤㻘 㻞㻜㻝㻠㼀㼔㼑 㻞㼚㼐㻦 㻭㼡㼓㻚 㻞㻥㻙㻟㻜㻘 㻞㻜㻝㻠㼀㼔㼑 㻟㼞㼐㻦 㻹㼍㼞㻚 㻜㻡㻙㻜㻤㻘 㻞㻜㻝㻡㼀㼔㼑 㻠㼠㼔㻦 㻶㼡㼘㻚 㻞㻝㻙㻞㻞㻘 㻞㻜㻝㻡㼀㼔㼑 㻡㼠㼔㻦 㻹㼍㼞㻚 㻜㻟㻙㻜㻢㻘 㻞㻜㻝㻢㼀㼔㼑 㻢㼠㼔㻦 㻿㼑㼜㻚 㻝㻞㻙㻝㻡㻘 㻞㻜㻝㻢㼀㼔㼑 㻣㼠㼔㻦 㻹㼍㼞㻚 㻜㻞㻙㻜㻡㻘 㻞㻜㻝㻣㼀㼔㼑 㻤㼠㼔㻦 㻿㼑㼜㻚 㻞㻢㻙㻞㻤㻘 㻞㻜㻝㻣㼀㼔㼑 㻥㼠㼔㻦 㻹㼍㼞㻚 㻜㻟㻙㻜㻡㻘 㻞㻜㻝㻤㼀㼔㼑 㻝㻜㼠㼔㻦 㻿㼑㼜㻚 㻞㻞㻙㻞㻠㻘㻞㻜㻝㻤㼀㼔㼑 㻝㻝㼠㼔 㻦 㻹㼍㼞㻚 㻜㻝㻙㻜㻟㻘 㻞㻜㻝㻥