Pan Africa News
Newsletterof theCommittee
Care
TheTheNewsletterof theCommittee
for forthetheCare
Conservation
Chimpanzees,
andandConservationof ofChimpanzees,
andandthethe
MahaleWildlife
Conservation
Society
MahaleWildlife
Conservation
Society

JUNE 2007
P. A. N. EDITORIAL STAFF

VOL. 14, NO. 1

Contents

Chief Editor:
Toshisada Nishida, Japan Monkey Centre, Japan
Deputy Chief Editors:
Kazuhiko Hosaka, Kamakura Women’s University, Japan
Michio Nakamura, Kyoto University, Japan<NOTE>
Laterality in Termite-Fishing by Fongoli Chimpanzees:
Preliminary Report

Associate Editors:
Christophe Boesch, Max-Planck Institute, Germany
Jane Goodall, Jane Goodall Institute, USA
Takayoshi Kano, Kyoto University, Japan
Tetsuro Matsuzawa, Kyoto University, Japan
William C. McGrew, University of Cambridge, UKJohn C. Mitani, University of Michigan, USA
Vernon Reynolds, Budongo Forest Project, UK
Yukimaru Sugiyama, Kyoto University, Japan
Richard W. Wrangham, Harvard University, USA<NOTE>
Evidence of Cultural Differences in Diet between Two
Neighboring Unit Groups of Chimpanzees in Mahale
Mountains National Park, Tanzania

Editorial Secretaries:
Noriko Itoh, Kyoto University, Japan
Koichiro Zamma, Japan Monkey Centre, Japan
Agumi Inaba, Japan Monkey Centre, Japan

Instructions to Authors:
Pan Africa News publishes articles, notes, reviews,
forums, news, essays, book reviews, letters to editor, andclassified ads on any aspect of conservation and research
regarding chimpanzees (Pan troglodytes) and bilias (Panpaniscus). Contributors are requested to write in English andthe papers except forums, reviews and essays should usually
be 1,500 words or less. Manuscripts should be submitted bye-mail to:
pan_ed@jinrui.zool.kyoto-u.ac.jp
Photos and figures, however, should be sent by air mailto: T. Nishida, Department of Zoology, Graduate School of
Science, Kyoto University, Kitashirakawa- Oiwakecho, Sakyo,
Kyoto, Japan.
Publication of the next issue will be December 2007.
Deadline for manuscripts is the end of November.

<NOTE>

Laterality in Termite-Fishing by
Fongoli Chimpanzees:
Preliminary Report
Paco Bertolani1, Clarissa Scholes1,
William C. McGrew1, Jill D. Pruetz2

Paco Bertolani, Clarissa Scholes, William C. McGrew& Jill D. Pruetz

Tetsuya Sakamaki, Michio Nakamura & Toshisada Nishida

1

3<NOTE>
Dermatophytosis of M group Chimpanzees, Mahale
Mountains, Tanzania
Toshisada Nishida, Shiho Fujita, Takahisa Matsusaka,
Masaki Shimada & Rashidi Kitopeni

5<NOTE>
Bonobos at the “Lola Ya Bonobo” Sanctuary in the
Democratic Republic of the Congo
Satoshi Hirata & Yasuko Tashiro

6<NOTE>
Snare Removal for Conservation of Chimpanzees in the
Kalinzu Forest Reserve, Uganda
Chie Hashimoto, Debby Cox & Takeshi Furuichi

8<NOTE>New Nut-Cracking Sites in Diecké Forest, Guinea: An
Overview of the Surveys
Susana Carvalho, Cláudia Sousa & Tetsuro Matsuzawa

11<NOTE>
Papaya Fruit Sharing in Wild Chimpanzees at Bossou,
GuineaGaku Ohashi

14

1. Leverhulme Centre for Human Evolutionary Studies, Dpt. ofBiological Anthropology, University of Cambridge, UK
2. Dpt. of Anthropology, Iowa State University, USA

INTRODUCTION
Many studies in both free-ranging and captive apes
have shown that some forms of laterality of hand function
occur in non-human primates1. However, true handedness(sensu McGrew and Marchant2), when most individuals

1

2

Pan Africa News, 14(1), June, 2007

show a skew in hand preference in the same direction

analysis was done only on individual sessions, which doacross different tasks, seems to be restricted to humans.

not equal events 2. Laterality was tested with the binomial

Other hominoids appear unlateralized in simpler tasks,

test for each individual who attained at least 6 sessions.

3such as reaching, picking up objects, and grooming , but

All tests are two-tailed, with alpha set at 0.05 (A further

they show hand preference for more complex tasks, such

10 chimpanzees showed termite-fishing but did not reach2, 4, 5as tool-using

6, 7

this criterion, which is the minimum number necessary toor elaborate food processing .
8

Laterality in termite-fishing has been studied only

show a statistic significance).
PB recorded which hand inserted and extracted theat Gombe, and the two published data-sets are congruent.
2, 9McGrew and Marchantreported that most (27 of 36)

chimpanzees showed an individualized hand preference

tool into and from the hole in the termite mound for eachinsertion event.

10

for right or left, as did Lonsdorf and Hopkins (16 of 17)
for termite-fishing in the same community. No other data

RESULTS

have been published for chimpanzee communities

Table 1 shows the data for 47 sessions ofelsewhere. This study asks if termite-fishing by Fongoli

termite-fishing shown by 5 males. Each session averaged

chimpanzees is lateralized, shows hand preference

9.1 data-points (range per individual: 7.3-10.6) and lasted(individuals are lateralized, but with no populationalan average of 45 minutes. Two individuals, BN and KL,

preference for either hand), or task specialization (all or

were exclusively lateralized to the left and rightmost individuals use the same hand).

respectively; YO was significantly right lateralized, andMM and SI were right-biased and left-biased respectively,
but not significantly so. Although the sample is very

METHODS
Fongoli, in southeastern Senegal, is a mosaic

small, no skew emerges in the direction of handsavanna, and the only hot, dry and open study site withpreference among these subjects. The results for11

habituated chimpanzees (see Pruetz, 2006 , for a
description of the habitat and the diet of Fongoli
chimpanzees).

The

only

published

study

of

instantaneous samples reflect those for sessions.
Table 1. Frequency of lateralized (L vs. R) termite-fishing by
sessions and instantaneous samples.

termite-fishing at Fongoli is based on indirect data12(Bogart and Pruetz, in prep.).
PB collected these data in April and May 2005,
after habituation of the first males of the Fongoli
community. The chimpanzees were observed for 543

Subject

Obs

LH
sessions

RH
sessions

LH inst.
samples

RH inst.
samplesp-value

BN

6

6

0

KL

8

0

8

MM

16

4

SI

8

7

Hand
preference

41

0

0.032

L

0

74

0.008

R

12

27

130

0.076

(R)

1

56

25

0.07

(L)

hours over 50 days, when termites were a major food

YO

9

1

8

14

52

0.04

R

source. PB collected data using scan sampling with a

Total

47

18

29

138

281

-

-

focal subject target on the subject’s activity and on theother individuals in sight, by instantaneous recording
13

DISCUSSION
Termite-fishing at Fongoli appears to be anevery 5 minutes .
A session of termite-fishing was a continuous

individually lateralized task, in which some of theperiod during which the individual ate termites. Sessionsindividuals specialize in using one hand, and some use

are separated by other activities (e.g. rest, groom, etc).

the other, but none is ambilateral. These results agree

During a session, change of posture or position andwith studies of Gombe chimpanzees3, 9, 10, despite thechange of tool occurred: thus a session contained several

sites being separated by thousands of kilometers on

2

bouts (sensu McGrew and Marchant ). The data-points

opposite sides of Africa. Extending data collection to thetaken in each session by instantaneous sampling cannot

rest of the Fongoli community is needed to validate thesebe considered independent observations, so statisticalpreliminary findings.

3

Pan Africa News, 14(1), June, 2007
Acknowledgments. We thank the Republic of Senegal andDepartment of Eaux et Forets for permission to work in
Senegal; D. Kante and M. Camara for assistance in the field. J.P.
was supported by grants from the National Geographic Society,
Center for the Study of Violence at Iowa State University (ISU),ISU Foreign Travel Grant, ISU Faculty Professional
Development Grant, and the American Society of Primatologists
Conservation Grant.

REFERENCES
1. McGrew WC, Marchant LF 1997. On the other hand:Current issues in and meta-analysis of the behavioral lateralityof hand function in nonhuman primates. Yearb Phys Anthropol
40: 201-232.
2. McGrew WC, Marchant LF 1996. On which side of theapes? Ethological study of laterality of hand use. In: Great Ape
Societies, McGrew WC, Marchant LF, Nishida T (eds),
Cambridge: Cambridge University Press, pp. 255-272.

<NOTE>

Evidence of Cultural Differencesin Diet between Two Neighboring
Unit Groups of Chimpanzees in
Mahale Mountains National Park,
Tanzania
Tetsuya Sakamaki1, Michio Nakamura2,
Toshisada Nishida3
1. Faculty of International Studies, Meiji Gakuin University, Japan
2. Graduate School of Science, Kyoto University, Japan
3. Japan Monkey Centre, Japan

Assessing behavioral diversity is key to investigating

3. Marchant LF, McGrew WC 1996. Laterality of limb
function in wild chimpanzees of Gombe National Park:
Comprehensive study of spontaneous activities. J Hum Evol 30:
427-443.

the existence of culture in wild chimpanzees and4. Boesch C 1991. Handedness in wild chimpanzees. Int J
Primatol 12: 541-558.

chimpanzees has come from comparisons of distantly

5. Sugiyama Y, Fushimi T, Sakura O, Matsuzawa T 1993.
Hand preference and tool use in wild chimpanzees. Primates 34:
151-159.
6. Byrne RW, Byrne JME 1993. Complex leaf-gathering skillsof mountain gorillas (Gorilla g. beringei): Variability andstandardization. Am J Primatol 31: 241-261.
7. Corp N, Byrne RW 2002. Leaf processing by wild
chimpanzees: Physically defended leaves reveal complex
manual skills. Ethology 108: 673-696.
8. Goodall J 1963. Feeding behaviour of wild chimpanzees. A
preliminary report. Symp Zool Soc Lond 10: 39-47.
9. McGrew WC, Marchant LF 1992. Chimpanzees, tools, andtermites: Hand preference or handedness? Curr Anthropol 33:
114-119.
10. Lonsdorf EV, Hopkins WD 2005. Wild chimpanzees show
population-level handedness for tool use. Proc Natl Acad Sci
USA 102: 12634-12638.
11. Pruetz JD 2006. Feeding ecology of savanna chimpanzees(Pan troglodytes verus) at Fongoli, Senegal. In: Feeding
Ecology in Apes and Other Primates: Ecological, Physical andSocial Aspects, Hohmann G, Robbins MM, Boesch C (eds),
New York: Cambridge University Press, pp. 161-182.

determining their ability to adapt to various environments1.
However, most evidence of behavioral diversity among
separated populations2. Although a few reports havesuggested differences in social customs between two
neighboring unit groups3, 4, these studies were retrospective
comparisons using photographs. Thus, a need exists toinvestigate whether any behavioral differences occur
between neighboring groups sharing a similar environment.
We report preliminary evidence of new feeding habits bychimpanzees that we are habituating, habits that differ
from those of a neighboring group in the Mahale
Mountains National Park, Tanzania.
A new unit group under habituation
Several different unit groups of chimpanzees live in
Mahale Mountains National Park. Intense studies of twoneighboring unit groups, in particular the K and M groups,
have been conducted since 19655. The K group was
declared extinct in the 1980s, leaving the M group as the
only habituated chimpanzees in the park6. However, one

12. McGrew WC, Pruetz JD, Fulton SJ 2005. Chimpanzees
use tools to harvest social insects at Fongoli, Senegal. Folia
Primatol 76: 222-226.

adult male of the K group, Limongo, was observed in the

13. Altmann J 1974. Observational study of behaviour:
sampling methods. Behaviour 49: 227-265.

the late 1990s, unhabituated chimpanzees were observed

former range of the K group in the 1980s and 1990s7. In
and heard occasionally in the former range of the K group8.
We started habituating these chimpanzees in 2005.
After

two

observational

periods(September

4

Pan Africa News, 14(1), June, 2007

2005–February 2006 and September 2006–December

suggesting that at least some chimpanzees of the Y group2006), we concluded that the group we were habituating

eat the young liana stems constantly and spit out thewas different from the former K group because we werewadges.

unable to positively confirm individuals from the former K
group, such as Limongo. Thus, we named this group the Y
group (representing the Miyako group). Judging fromsome indirect evidence so far, such as vocalizations andfood remnants, we assumed that the Y group occupied
about the half of the former K-group range. Although the
maximum number of individuals directly observed at any
one time was eight, we heard loud calls exchanged fromother parties. Based on these calls, we estimated a groupsize of about 50 chimpanzees.
Evidence of feeding on wood-boring ants, Camponotus
brutusFig. 1. A wadge of young stems of Cissus oliveri liana, chewed
and spit out presumably by a Y group chimpanzee

In October 2005, we noticed Camponotus brutus antsin the feces of Y-group chimpanzees. These large reddish
ants inhabit the ranges of both the M and Y group;
however, the M group chimpanzees only rarely eat them.
The ants are extremely aggressive and powerful, making it
difficult to catch and eat them without getting hurt. The
chimpanzees typically eat a related species, C. vividus,
using a strip of bark fiber or other item as a tool to fishthem out of their tunnels9.
In the 2006 season, however, we identified C. brutus
ants in at least 21 of 41 fecal samples, suggesting that theants are a common food item for Y-group chimpanzees.
We have not yet confirmed their feeding techniques, but
given the aggressive nature of these ants and the large
number consumed, the chimpanzees may use some kind oftool to harvest these ants.
Evidence of eating young stems of liana, Cissus oliveri
In September 2006, we found some unknown fibrouswadges from chimpanzees. We identified them in the field
as a type of vine or liana, but could not specify the species.
Later, we found half-chewed leaves that were similar to thewadges (Fig. 1). These were young stems of Cissus oliveri
liana, which are rarely eaten by chimpanzees of the M andK groups10, even though the plant is abundant throughout

It is only natural to presume that potential foodspecies are not very different among groups, consideringthat the range of the Y group is almost identical to the halfof the former K-group range and is adjacent to and partly
overlaps M group’s range. We confirmed this with data on
the diet of the K and M groups10. Therefore, our findings
strongly suggest that the Y group has developed a different
dietary preference that cannot simply be explained by theavailability of food species. Moreover, the relative
abundance of these new food items throughout the range
indicates that they are consumed in common by Y groupmembers.
In future work, we need to confirm (via direct
observation) the techniques with which the Y-groupchimpanzees harvest and eat these new food items.
Although we are still in the process of habituating andidentifying individual chimpanzees of this group, we
expecttofindbehavioral

diversity

acrossneighboring unit groups.
Acknowledgments. This research was financially supported by aGrant for the Biodiversity Research of the 21st Century COE(A14) and MEXT (16255007 and 19255008 to TN and 16770186
to MN). We thank the COSTECH, TAWIRI, TANAPA, MMNP,
and MMWRC for permission to conduct the fieldwork and forlogistical support.

most of their ranges. Throughout the remainder of thestudy period, we found 23 more wadges of C. oliveri,

more

REFERENCES

5

Pan Africa News, 14(1), June, 2007
1.
Boesch C, Hohmann G, Marchant LF (eds) 2002.
Behavioural diversity in chimpanzees and bonobos. Cambridge
University Press, Cambridge, UK.
2. Whiten A, Goodall J, McGrew WC, Nishida T, Reynolds V,
Sugiyama Y, Tutin CEG, Wrangham RW, Boesch C 1999.
Cultures in chimpanzees. Nature 399: 682–685.
3. McGrew WC, Marchant LF, Scott SE, Tutin CEG 2001.
Intergroup differences in a social custom of wild chimpanzees:
The grooming hand-clasp of the Mahale Mountains. Curr
Anthropol 42: 148–153.
4. Nakamura M, Uehara S 2004. Proximate factors of two typesof grooming-hand-clasp in Mahale chimpanzees: Implication for
chimpanzee social custom. Curr Anthropol 45: 108–114.
5. Nishida T (ed) 1990. The chimpanzees of the Mahale
Mountains: Sexual and life history Strategies. University ofTokyo Press, Tokyo.
6. Nishida T, Hiraiwa-Hasegawa M, Hasegawa T, Takahata Y
1985. Group extinction and female transfer in wild chimpanzees
in the Mahale National Park, Tanzania. Z Tierpsychol 67:
284–301.
7. Uehara S, Nishida T, Takasaki H, Kitopeni R, Kasagula MB,
Norikoshi K, Tsukahara T, Nyundo R, Hamai M 1994. A lone
male chimpanzee in the wild: The survivor of a disintegrated
unit-group. Primates 35: 275–281.
8. Uehara S 2002. Evidence of the leaf-clipping behavior bychimpanzee of an unhabituated group at Mahale. Pan Afr News 9:
3–4.
9. Nishida T, Hiraiwa M 1982. Natural history of a tool-using
behavior by wild chimpanzees in feeding upon wood-boring ants.
J Hum Evol 11: 73–99.
10. Nishida T, Uehara S 1983. Natural diet of chimpanzees(Pan troglodytes schweinfurthii): Long-term record from the
Mahale Mountains, Tanzania. Afr Study Monogr 3: 109–130.

40 years1. We noticed a skin disease developing amongsome members of M group in 2001 and 2002.
Table 1. List of chimpanzees that suffered from dermatophytosis
Individual

Age & SexFirst daydermatophytosis
observed

Last daydermatophytosis
observed

Minimum
duration(Days) ofsuffering

Caesarinfant male

May 10, 2001

May 29, 2001

20

Ruby

adult female

May 21, 2001

Jun. 3, 2001

14

Rubiconinfant female

Jun. 3, 2001

Jul. 2, 2001

29

Michiojuvenile maleAug.30, 2001

Oct. 3, 2001

35

Alofu

adult male

Sep. 9, 2001

Oct. 3, 2001

25

Darwinadolescent male

Sep.16, 2001

Oct. 9, 2001

24

Jiddahjuvenile female

Sep.16, 2001

Oct. 2, 2001

17

Miya

adult female

Sep.21, 2001

Sep.30, 2001

10

Cadmusadolescent male

Sep.25, 2001

Oct. 3, 2001

9

Aqua

infant female

Sep.29, 2001

Sep.30, 2001

2

Ivana

juvenile female

Nov. 8, 2001

Nov.21, 2001

14

Cynthia

adult female

May 18, 2002

May 18, 2002

1

Xmas

juvenile male

Oct. 4, 2002

Oct. 4, 2002

1

RK observed M group chimpanzees for the wholeyears of 2001 and 2002, SF from June to October 2001,
TM from September 2000 to June 2001, TN from
September to October 2001 and from September toNovember 2002, and MS from September 2001 to October
2002. We recorded any observable injuries, disease
symptoms such as coughing, and abnormalities of the skin,faces and eyes.
On May 10, 2001, we first noticed that an infant<NOTE>

Dermatophytosis of M group
Chimpanzees, Mahale
Mountains, Tanzania
Toshisada Nishida1, Shiho Fujita2,
Takahisa Matsusaka1, Masaki Shimada3,
Rashidi Kitopeni4

male, Caesar, had a whitish nose and left cheek; it looked
as if white chalk had been applied to the part. On July 2, afemale infant, Rubicon, had a whitish face. On September
9, an adult male, Alofu, had a whitish upper lip. In
September, as many as 7 individuals at once showed thisskin disease.
Table 1 shows the first and last observed dates of thedisease in 13 individuals. Its occurrence biased to seasons

1. Japan Monkey Centre, Japan

with high humidity might suggest a fungal infection, so

2. Yamaguchi University, Japanthat the disease may be some form of dermatophytosis.

3. University of Shiga Prefecture, Japan
4. Mahale Mountains Chimpanzee Research Project, Tanzania

Victims were from all age-sex classes and no conspicuous
age or sex differences were found (1 of 8 adult males, 3 of

The chimpanzees of the Kasoje area in the Mahale

20 adult females, 2 of 5 adolescent males, 0 of 2 adolescent

Mountains National Park have been studied for more thanfemales, 2 of 2 juvenile males, 2 of 3 juvenile females, 1 of

6

Pan Africa News, 14(1), June, 2007

3 infant males, 2 of 10 infant females).
These

12

dermatophytosis

chimpanzees

had

on

in

their

faces,

particular, upper lip, nose, cheek, andsupraorbital ridge (Fig.1). The exception
was Michio, who had a white mark on theback of his hand, as well as a white face.
Although we could not always check allindividuals during the study periods, it
was likely that the skin disease lasted
several weeks (minimum, median = 14days) for each victim and the longest span
was 35 days. The infectious nature of thedisease was inferred from the victims
being concentrated to the short period of
September to October, 2001 and from theclose relationships among at least someFig. 1. Dermatophytosis on the faces of the Mahale M-group chimpanzees. See thetext for details.

victims: for example, Miya is the mother of Michio, andRuby is the mother of Rubicon. Roy2 described a fungal
disease from the chimpanzees of Gombe, but not this typeof skin disease. Bill Wallauer’s3 recent blog shows aphotograph of a Gombe chimpanzee, Faustino, who<NOTE>

Bonobos at the “Lola Ya Bonobo”
Sanctuary in the Democratic
Republic of the Congo

appeared to show the same skin disease as described here.
Neither victims nor the others showed any attention
to the infected parts of the body; the victims did not scratch
the whitish parts of their faces. Later, all individuals
recovered their normal dark faces without experiencing
any apparent health problems (Cf. Hubbard4).
Acknowledgments. The research was funded by MEXT Scientific
Research Fund A1 (#12375003 & 16255007 to TN) and Global
Environment Research Fund of the Ministry of Environment(F-061 to TN). We thank Bill McGrew for useful comments.

REFERENCES

Satoshi Hirata, Yasuko Tashiro
Great Ape Research Institute,Hayashibara Biochemical Laboratories, Inc., Japan

Great apes face extinction from threats mainly causedby human activities such as logging in habitat areas,
bush-meat hunting, and pet trade. Many African countries
have domestic laws prohibiting the capture and trade ofendangered great apes from the wild. However, theenforcement of these laws is difficult without addressing
how to care for illegally owned animals. To address the

1. Nishida T (ed) 1990. The Chimpanzees of the Mahale
Mountains. Tokyo: The University of Tokyo Press.

problem, various “sanctuaries” have been established to

2. Roy AD 1974. Rhinophycomycosis entomophthorae occurringin a chimpanzee in the wild in East Africa: further report.
American Journal of Tropical Medicine and Hygiene, 23:935.

example, 13 chimpanzee sanctuaries in 11 African

3. Wallauer W 2007. The Jane Goodall Institute’s Gombe
Chimpanzee Blog

However, there is only one bonobo sanctuary in the world,(http://www.janegoodall.org/Gombe-Chimp-Blog/detail.asp?Entry_ID=44)

4. Hubbard GB 2001. Nonhuman primate dermatology.
Veterinary Clinics of North America: Exotic Animal Practice,
4:573-583.

provide homes for confiscated great ape orphans. For
countries now house over 600 chimpanzee orphans1.
the Lola Ya Bonobo sanctuary, located approximately 25
km southwest of Kinshasa, the capital of the Democratic
Republic of the Congo, and directed by Ms. Claudine
André2.

7

Pan Africa News, 14(1), June, 2007
The sanctuary was created bytheir infants, and youngsters who

Les Amis des Animaux au Congo

had been “weaned” from human(AAC; “Friends of Animals in

substitute mothers lived together.

Congo”), a Congolese organization

The first group, consisting of 15

3created in 1994 by Ms André . She

individuals, used one of the largeorganized the AAC to support

enclosures. The second group,

animals in the Kinshasa Zoo. Soonconsisting of 13 individuals, used

afterwards, an illegally owned

the other large enclosure. The third

bonobowasconfiscatedincollaboration with the government,group, consisting of 11 individuals,

Fig. 1. A bonobo walking in water.

used the mid-sized enclosure. The

and the AAC began to care for young orphan bonobos. In

last group consisted of 6 individuals who were kept in the

1998, the AAC moved the bonobos to the grounds of thesmall enclosure.

a

All members of the mixed-age groups were kept innursery-sanctuary, which continued to receive illegallyindoor quarters during the night. In the morning (aroundcaptured bonobos. In 2002, the sanctuary was moved to its

0600 hr), staff members gave them water supplemented

current location and changed its name to Lola Ya Bonobo;

with minerals and sugar and visually inspected the healthmeanwhile AAC also changed its name to Les Amis desof the bonobos. The groups were then released into their

Bonobos du Congo (ABC; “Friends of Bonobos inrespective enclosures at around 0630 hr after the safety ofCongo”). Lola Ya Bonobo now occupies about 30 ha ofthe enclosures was ensured. Occasionally, the bonobos

land previously owned by a French businessman. This area

received external applications of anti-parasite medicine

is divided into five enclosures for the bonobos: two

before being released into the enclosures. At other times,

large-sized, one middle-sized, and two small-sizedwhile the bonobos were still in their nighttime quarters,

enclosures. The enclosures, surrounded by electric fences,

workers mowed the grass near the electric fence to prevent

include 20 m tall forest areas, swampy areas, and a small

a short circuit. During the day, staff members gave theriver; thus, the environment is complex and similar tobonobos various types of fruits, vegetables, sugarcane,natural bonobo habitat.

peanuts, breads, eggs, and milk. In addition to caring forAmericanschoolin

Kinshasa

andcreatedIn February 2007, we had the opportunity to visit

the bonobos, the sanctuary sets a high priority on

Lola Ya Bonobo and stay for 3 weeks. At the time of oureducation4. Educational tours for Congolese children took

stay, 53 bonobos lived there, including four infants born inplace during our visit, and the children eagerly observed

the sanctuary. The following is a summary of our

the bonobos and attended the talks given by the staff.

observations during the 3 weeks, which we hope will
illustrate daily life at the sanctuary.

The composition of bonobo members within a groupis decided by taking into consideration their compatibility

Among the 53 bonobos, eight were 3–5 year olds

with each other. Young orphans who are dependent on

who were dependent on human substitute mothers. Therehuman mothers will be integrated into a social group andwere about 20 local people working for the sanctuary. Of

begin to learn necessary skills such as foraging and danger

these, four worked as substitute mothers. They took care of

avoidance from conspecific group members. We had thethe young bonobos from morning to evening, cleaning

opportunity to observe the integration of groups during ourthem, giving them milk and other foods, and letting themstay. Three of the six individuals in the small enclosure

play in a play yard. In the evening, the bonobos were

were young individuals (approx. 4–5 years of age) who

brought to cages to spend the night.

had just finished their time of care by human mothers.

At the beginning of our stay, the remaining 45

Three subadult (approx. 8–9 years of age), good-tempered

individuals lived in four mixed-age groups in which adults,

individuals were chosen from the first mixed-age group,8

Pan Africa News, 14(1), June, 2007

and these three older and three younger bonobos were

behaviors that have not been observed in their wildallowed to spend time with each other in the small

counterparts such as stone-tool use to crack open oil palm

enclosure. On 12 February 2007, four individuals from thenuts, the use of plant materials to drink water, and the use

first mixed-age group were moved to the group in theof probing sticks. One conspicuous feature was theirsmall-sized enclosure as the next step in the integration offamiliarity with water. They did not literally swim, but

the young individuals. Two of these four individuals werewould enter the water to just below their shoulders (Fig. 1).

adult females who were expected to protect the young

These examples show the flexibility of their behavior andindividuals after integration; another was a juvenile, who

the great influence of post-natal experience for bonobos.

was included as a possible playmate; and the last was aThe

sanctuaryis

now

considering

possiblemale over 10 years old. The bonobos emitted many

reintroductions of bonobos into the natural habitat as

vocalizations and engaged in affinitive interactions such as

another tool for managing wild bonobo populations4. Thisembracing and GG-contact during their first encounter; thelone bonobo sanctuary thus plays important roles in

excitement ceased within approximately 1 hour, without

conservation, education, and research on this endangered

any aggressive interactions. On the following day, theclosest relative of humans.

members of the small-sized enclosure, including the three
young individuals, were released into the large enclosure tojoin the remaining 10 members of the first mixed-age
group. An adult male performed display behavior and ran
around the newcomers, but did not show any aggressive
physical contact. All of the members gathered around thenewcomers and engaged in numerous vocal andsocio-sexual exchanges. Most of the members thenclimbed a large, 20 m tall tree and continued to make vocal
exchanges; the event ended peacefully after approximately
1 hour.
The integration of the second and third mixed-age
groups was also carried out over the course of the next few
days. The reactions of bonobos during their firstencounters were similar to those described above. They
engaged in many vocal and socio-sexual exchanges, andAcknowledgments. Our study was financially supported by
MEXT (18700266 to Satoshi Hirata and 17255005 to Takeshi
Furuichi). We thank Claudine André for giving us the opportunity
to stay at Lola Ya Bonobo and all of the other staff for theirsupport during our stay. We also thank Brian Hare for his
generous guidance concerning our visit.

REFERENCES
1. Kabasawa A 2006. The current status of chimpanzee
sanctuaries: how do we deal with “chimpanzees with no place togo” (in Japanese). Ecosophia 17: 88–103.
2. André C 2006. Une tendresse sauvage (in French).
Calmann-Lévy, Paris.
3. Ihobe H, Idani G, Tashiro Y 2006. Current status of Lola Ya
Bonobo (in Japanese). Primate Res 22: 37–41.
4. André C, Kamate C, Mbonzo P, Morel D, Hare B in press. The
conservation value of Lola Ya Bonobo sanctuary. In: Thebonobos: behavior, ecology, and conservation, Thompson J,
Furuichi T (eds), New York: Springer.

the integration ended without harmful interactions after
approximately 1 hour. After a series of integrations andrecombinations, two new mixed-age groups had formed by
the end of our stay: the first group consisted of 19
individuals (10 males and 9 females, 0–21 years old),
excluding a female separated for preparation for delivery
and a male selected as her cagemate who were originallymembers of this group; the second group consisted of 24
individuals (18 males and 6 females, 1–23 years old).
While in their enclosures, the bonobos engaged ingrooming, solo and social play, GG-contact, and otheractivities typical of their species. They also displayed some

<NOTE>

Snare Removal for Conservationof Chimpanzees in the Kalinzu
Forest Reserve, Uganda
Chie Hashimoto1, Debby Cox 2, Takeshi Furuichi3
1. Primate Research Institute, Kyoto University, Japan
2. Jane Goodall Institute Uganda
3. Faculty of International Studies, Meiji-Gakuin University, Japan

9

Pan Africa News, 14(1), June, 2007
INTRODUCTION
While hunting of primates, particularly chimpanzees,
is not common in Uganda, hunting of other forest
mammals, such as duikers and bushpigs is known to occur.
This hunting has an indirect impact on chimpanzees andother primates as most of the hunting in Uganda is done bysetting snares or jaw traps. As chimpanzees walk throughthe forest, their hands or feet may become trapped in thesnare made of metal wire1. In two of the forests in Uganda(Kibale and Budongo), researchers have observed up to
25% of chimpanzees with snare-related injuries1, 2, 3. In the
Kalinzu Forest, half of the adult males in the M group hadsnare injuries (Fig. 2)4.
In order to address such a critical situation, we started
the snare removal program with the support of Disney
Wildlife Conservation Fund, and continued the programwith the support of Japan Ministry of Environment Global

Fig. 2. An adult male who lost his foot by snare injury.

the beginning of May. Though we are still carrying out thisprogram, here we report the results from May 2005 to June
2006.

METHODS

Environment Research Fund. We started recruiting and

The Kalinzu Forest Reserve, which covers an area of

training staff in April 2005, and started the program from137 km2, is a part of Uganda’s largest forest block where
ecological survey of wild chimpanzees has been
undertaken since 19925. There inhabits at least four groups(communities) of chimpanzees (Fig. 1), with an estimatedpopulation of 230 individuals (1.67 chimpanzees per km2)6.
The main study group, called the M group, comprising of
18 adult males, 24 adult females, and 26 immatureindividuals in January 2007.
We employed four persons to form two teams for thesnare removal. Each consists of one ex-hunter who has a
deep knowledge on the geography and technique for
hunting in this area, and another person who graduatedfrom primary school and is used to speaking and writing
English. In order to raise local communities’ awareness of
this program, all persons were recruited from the villages
adjacent to the Kalinzu Forest Reserve.
Kalinzu Forest Reserve consists of 43 compartments(Fig. 1). The northern area of the reserve is better
conserved and has higher population of chimpanzees,
monkeys and other animals. We have trail systems forresearch in the northeast: compartments 25 to 30 and 36 to

Fig.

1.

Forestcompartments

andranging

chimpanzee groups in the Kalinzu Forest Reserve.

areas

of

40. Two habituated groups of chimpanzees inhabit this area,

10

Pan Africa News, 14(1), June, 2007

and we mainly targeted the northern area in the snare

employment produced by the research and conservation

removal program in this period.

activities in the villages adjacent to these compartments.

The two teams patrolled different parts of this area

On the other hand, snares were most frequently foundfrom Monday to Saturday. When they found a lot of snares

in compartment 40, 42, and 43, though we visited there

in some areas, they revisit these areas more intensively

once a month. There are newly established villages to thethan others. It was difficult to cover compartments 43, 42

north of the compartment 43. Because those villages are

and 40, due to lack of transportation for the teams. They

located far from the main road, people living there are notwalked through these compartments once a month fromwell

the northern edge of the reserve back to the main study

governmental organization which is responsible forarea.

management of Ugandan Forest Reserves. In addition,

controlledbyNational

Forestry

Authority,

those people have least opportunity to receive benefit fromRESULTS AND DISCUSSION

research, ecotourism, and conservation activities. How to

From May 2005 to June 2006, we found andcontrol the illegal activities in this area remains the biggest

removed 1022 snares and 136 live traps (Table 1). There

problem for the conservation of chimpanzees in thewere two types of snares: one made of nylon strings andKalinzu Forest Reserve.

one made of metal wire. The former was mainly used for

For one year since the beginning of the program,trapping guinea fowls, and the latter was used for trapping

there had been no tendency of decrease in the number ofbushpigs and duikers and both are harmful to thesnares removed. Although the slight increase might be a

chimpanzees1, 7.

result of the increased skill in finding snares, we may need

Table 1. Number of snares found in the Kalinzu Forest Reserve.
Year

Month

SnaresLive Traps

2005

May

71

6

Jun

53

6

2006

Jul

77

0

Aug

27

0

Sep

54

14

Oct

40

4

Nov

59

27

Dec

157

3

Jan

98

6

Feb

115

35

Mar

152

24

Apr

61

18

May

105

11

Jun

68

17

Total

1137

171

Total per month

81.2

12.2

* Though the snare removal program supported by Animal
Kingdom was terminated in April, the program is being continued
by the Kalinzu Forest Project.

Among the areas where we made daily patrols, snares
were frequently found in compartments 26, 27, 28, and 40
that are close to villages and the tea estate. However, not somany snares were found in compartments 37 and 38.
Though we need to wait for further studies, we expect thatthe lower number of snares in these compartments is theresults of our effort for conservation education andto continue this program for at least a few more years toevaluate its effect. Furthermore, we may need to develop
conservation program, such as education program, at thevillages adjacent to the compartments where snares weremost frequently found.
Acknowledgments. We thank staff of the Jane Goodall Institute,
the Kalinzu Forest Project, and the National Forestry Authority
for technical support. This study was financially supported by
the Disney Wildlife Conservation Fund and Japan Ministry ofEnvironment Global Environment Research Fund (#F-061 to T.
Nishida).

REFERENCES
1. Waller JC, Reynolds V. 2001. Limb injuries resulting fromsnares and traps in chimpanzees (Pan troglodytes schweinfurthii)of the Budongo Forest, Uganda. Primates 42:135-139.
2. Edroma E, Rosen N, Miller P (eds) 1997. Conserving the
Chimpanzees of Uganda: Population and Habitat Viability
Analysis for Pan troglodytes schweinfurthii. IUCN/SSC
Conservation Breeding Specialist Group, Apple Valley, MN.
3. Muller M 2000. Knuckle-Walking Wounded. Natural History
10/00:44-46.
4. Hashimoto C 1999. Snare injuries of chimpanzees in the
Kalinzu Forest, Uganda. Pan Afr News 6:20-22.
5. Hashimoto C 1995. Population census of the Chimpanzees in
the Kalinzu Forest, Uganda: Comparison Between Methods with
Nest Counts. Primates 36:477-488.

11

Pan Africa News, 14(1), June, 2007
6. Plumptre AJ, Cox D 2006. Counting primates for conservation:
primate surveys in Uganda. Primates 47:65–73.
7. Stokes EJ, Byrne RW 2006. Effect of snare injuries on the
fig-feeding behavior of chimpanzees of the Budongo Forest,
Uganda: behavioral adaptations and long-term implications. In:
Primates of western Uganda, Newton-Fisher NE, Notman H,
Paterson JD Reynolds V (eds), New York: Springer, pp.281-297.

important world spot for the biodiversity. These forests
survived after the retraction and division of Pleistocene
forests, partly due to the ice age that dried the climate
cyclically4.
This forest, Fôret Classée de Diecké, is a protectedarea of 600 km2, located in the South-eastern part ofGuinea, 50 km west from Bossou, bordering Liberia on thesouth. The reserve is controlled by the Centre Forestier de<NOTE>

N`zerekoré, and Diecké biodiversity is managed by a

New Nut-Cracking Sites inDiecké Forest, Guinea: An
Overview of the Surveys

German project located at the Direction Generale de1and organized the forest in thirty parcels with three levels

Centres Forestiers, also in N`zerekoré. This project,
entitled

PROGERFOR,

focusesonconservation/

reforestation; damage minimization; regional development,

2

Susana Carvalho , Cláudia Sousa ,Tetsuro Matsuzawa3of protection.

1. Dpt. Anthropology, Faculty of Sciences and Technology,
University of Coimbra, Portugal

Diecké is characterized by a large number of small
hills, averaging 350 m of altitude (the highest point being

2. Dpt. Anthropology, Faculty of Social and Human Sciences, New
University of Lisbon, PortugalMont Jna, with 800 m). Two main rivers cross the forest

3. Primate Research Institute, Kyoto University, Japanrunning south: Nyé on the west side, and Gbin, divided in
Gbin and Gbin-bé, on the east side. Secondary waterlines

INTRODUCTION

are abundant in the forest creating swamps. In this part of

We report the results of the 2006 KUPRI (Kyoto
University Primate Research Institute) surveys in Diecké
forest, Guinea, Western Africa.

the country, the dry season is very short, from December orJanuary to March or April.
Geologically, Diecké is part of the West African

Research in the Diecké forest by KUPRI started in

craton with Precambrian rocks, essentially igneous and1999 with a preliminary survey1. This
continued in 2000, with the survey of Nonahand Yossono areas2. The main objective wasto find a good research site that would allow
for comparative studies between different
communities of wild chimpanzees. It has
been acknowledged, since the first fieldseason, that Diecké chimpanzees crack open
nuts (Panda oleosa and Coula edulis) usingstone and root anvils and stone hammers.
Nut-cracking behaviour had not been
observed directly until today and there arevery few reports on chimpanzee’s direct
observation2, 3.
The Diecké forest is part of the
Guinean Forests and integrates the region

Fig. 1. One of the satellite views of Diecké Forest covering the surveyed areas incomprised by lowland forests, being one

Nonah area and Korohouan area (Credits: S. Carvalho / PRI-Kyoto, 2006).

12

Pan Africa News, 14(1), June, 2007

metamorphic5. Concerning raw material availability in thehours walking distance from Nonah to NNE direction.

primary forest zone, i.e. stone types and nut species,

Here it was recorded a total of seventeen tools: elevenenormous granite outcrops were observed and movablehammers and six outcropping anvils.

quartz stones were found near waterlines.

In March another survey was undertaken over an

Nut species of Panda oleosa tree and Coula edulis

eight day period. The SB1 place had been visited bytree are inside the integral protected area (per. obs.), as

chimpanzees, allowing the recording of tool use. The base6

these are species from mature forests . Elaies guineensis iscamp was set to Northeast from SB1, crossing Lilaya

not present in the primary forest; however it is possible towatercourse.

locate natural, or human planted, Elaeis in the peripheral

During this trip another nut-cracking place (SB2) of
Panda oleosa nuts, was identified, located east of SB1.

area.

Seven tools were recorded: four hammers and three
The etho-archaeological surveys
In 2006, chimpanzee research continued in this area,

outcropping anvils. Raw materials were, preferentially,
granite and quartz (Table 1).

combining primatological and archaeological methods,

Survey continued in an easterly direction, passing thewhile new zones were surveyed. The KUPRI work team,

Nyé River (Fig.1), four hours walking from the base camp.

accompanied by local guides, carried out three survey

In this area, mature fruits of Monkey orange or Elephantincursions between January and April 2006 and set twoorange tree (Strychnos spinosa) were observed, as well as

temporary camps in the core of the forest, one departing

Parinari excelsa fruits.

from Nonah village (7º 31`50.9``N; 9º 04`27.7``W) andDuring another survey, the Kolpégue watercourse (7ºanother from Korohouan village (7º 26`09.6``N; 8º34`31, 4``N; 9º 01`43, 1``W) was crossed to arrive at the59`22.2``W) (Fig. 1).

Taffa river area (7º 34`26, 5``N; 9º 00`49, 4``W), five

A selected survey was performed searching for nut
trees. Once it was apparent that nut-cracking places werehours walking from the base camp. In Taffa area one
chimpanzee sleeping trace was recorded-an old bed.

located near water lines, the survey was directed to theseIn April, one survey was carried out over a period ofpoints. When one nut-cracking site was found, a radio

seven days. The goal was to survey the Korohouan area.

centric survey was attempted, 1km around the nut-cracking

Moving east, we crossed the Gole watercourse (7º 26`18,

area. As a result, six nut-cracking sites were recorded, four8``N; 8º 58`41, 3``W), Melu watercourse (7º 27`03, 5``N;of them previously unknown.

8º 57`20, 8``W), Gbin river (7º 27`08, 3``N; 8º 57`12,

A pilot survey was carried out in January, for three

5``W), Dené watercourse and Sakai watercourse. On thedays, departing from Nonah village. The aim was to collectright border of Sakai watercourse, a new nut-cracking site

information and to prepare local work teams to enter the(SB 3) of Panda oleosa, was found (Table 1). This site wasforest. During this survey, a nut-cracking site of Panda

abandoned some years before and showed a large

oleosa nuts was detected (SB1) (Table 1). Originallyarchaeological assemblage. Visible hammers were often

discovered by Matsuzawa1, SB1 is located around a threefractured and site characteristics were similar to SB1 andTable 1. Location of the six nut-cracking sites recorded during 2006 field season in Diecké
forest. The grey colour indicates the new sites.

SB2. A total of forty tools wererecorded:one

hammer,eight

outcropping anvils, and thirty-one
fragments of hammers.
The base camp was set on theleft border of Gbin-bé river (7º28`14, 6``N; 8º 54`33, 6``W). Freshtraces of chimpanzee feeding of

13

Pan Africa News, 14(1), June, 2007
Landolphyaowarensis

wereobserved in the base camp area.
Following to NNE, two new
nut-cracking sites of Coula edulis
were found (SB4, SB5) (Table 1).
These sites had been used about one
month before. In SB4 a total of six
tools were recorded: three hammers
and three outcropping anvils. In SB5eleven tools were detected: eighthammers and three outcropping
anvils. It was also observed thatplenty of Cola cordifolia fruits had
been eaten in this area.
The last survey departed from
Korohouan to the Southwest, outside

Fig. 2. The SB 6 site in the Kourouhan area (Mont Medou). Large quartz hammers to open

the Diecké protected area. The Mont

crack the Panda nuts were found on the top of the outcropping anvil. Refitting of stones

Medou area - a remaining fragment

was also detected.

of primary forest - was surveyed. A
new nut-cracking place (SB6) of Panda oleosa nuts wasfound on the right margin of the Kpayan River (Table 1).
Here, a total of ten tools were recorded: six hammers andfour anvils. Two hammers were fractured in several pieces(Fig. 2). Following the Kpayan River we reached the top ofa valley where 25 beds were (7º 24`07, 2``N; 8º 59`03,
4``W).
Chimpanzee feeding, sleeping and travel traces were
found in the core and periphery areas of Diecké forest; thelatter provided banana plantations and other agricultural
fields as alternative food resources. Future surveys will
determine the size and the home range of the possible
different groups. We further expect to continue thesymbiotic approach between archaeology and primatology
in order to understand some of the older chimpanzee
nut-cracking sites regarding technological and typologicalvariance among chimpanzee groups.
Acknowledgments. We thank the Direction National de laRecherche Scientifique et Technique, République de Guinée, and
Dr. Kourouma Makan, Director of the Institut de Recherche
Environnemental de Bossou for permission to conduct field workat Diecké. We also thank Dr. Werner Grimmelman and Progerfor.
The research was supported by Grants-in-Aid for scientificresearch from the Ministry of Education, Science, Sports, andCultureofJapan:
MEXT-16002001,
JSPS-HOPE,
JSPS-21COE-Kyoto-Biodiversity, and Global Environment
Research Fund (F-061 to T. Nishida) of the Ministry ofEnvironment, Japan. We thank B. Zogbila, H. Gbelegbe, G. Doré,
P. Goumy, J. M. Kolié, J. Malamu, L. Tokpa, A. Kbokmo, C.
Koti, O. Mamy, for the field support. We are grateful to Francisco
Almeida for comments on the manuscript.

REFERENCES
1. Matsuzawa T, Takemoto H, Hayakawa S, Shimada M 1999.
Diecké Forest in Guinea. Pan Afr News 6: 10-11.
2. Humle T, Matsuzawa T 2001. Behavioural diversity among thewild chimpanzee populations of Bossou and neighbouring areas,
Guinea and Cote d`Ivoire, West Africa: A preliminary report.
Folia Primatol 72(2):57-68.
3. Kormos R, Humle T, Brugière D, Fleury-Brugière M,
Matsuzawa T, Sugiyama S, Carter J, Diallo MS, Sagno C,
Tounkara EO 2003. The Republic of Guinea In: Status Survey andConservation Action Plan West African Chimpanzees, Kormos, R,
Boesch C, Bakarr MI, Butynski TM (eds), IUCN/SSC Primate
Specialist Group. Published by IUCN, Gland, Switzerland andCambridge. UK.
4. Caldecott J, Kapos V 2005. Great ape habitats: tropical moist
forests of the old world. In: World atlas of great apes and theirconservation, Caldecott J, Miles L (eds), California: University ofCalifornia Press.
5. Clark JD 1967. The atlas of african prehistory. Chicago &London: The University of Chicago Press.
6. White L, Abernethy K 1996. Guide de la vegetation de lareserve de la Lopé. Écofac Gabon. Multipress Gabon. Gabon.

14

Pan Africa News, 14(1), June, 2007

<NOTE>

Papaya Fruit Sharing in Wild
Chimpanzees at Bossou, Guinea
Gaku Ohashi
Primate Research Institute, Kyoto University, JapanINTRODUCTION
Food sharing between non-kin of wild chimpanzees
has been reported since Goodall’s initial study in 19601.
Chimpanzees are often reported to share meat2, 3, 4, 5, 6. On

Fig. 1. Male chimpanzees came back from the village withthe other hand, plant food sharing are rarely reported7, 8, 9, 10,

papaya fruits.

except for studies from Gombe and Mahale where

METHODS

chimpanzees were formerly provisioned for research and

The subjects of this study were wild chimpanzees atobserved sharing sugar canes and bananas11, 12.

Bossou, Guinea18, 19. The home range size of this group is

Most studies discuss food sharing with reference toabout 30 km2, but the chimpanzees mainly use 4 km2 ofmaintenance of alpha status, social relationships andforest around the village of Bossou. Because their homecooperative hunting2, 3, 5, 6, 13, 14, 15. Some studies suggest arange is close to human settlements20, the chimpanzees

relationship between food sharing and female choice, but

often enter the village to raid crops (Fig. 1).

this claim lacks sufficient evidence6, 16 and recent studies

Observations were conducted from July 2002 tohave tried to refute it with substantial data13, 17.

March 2003 (Period 1) and from April to September 2004

I observed the cases of papaya fruit sharing by wild

(Period 2). Group size fluctuated between 13 and 19 during

chimpanzees at Bossou Guinea. The papaya is the largest

these periods. The number of adult or sub-adult males was

fruit that Bossou chimpanzees can obtain. Here I discuss

3. The number of cycling females fluctuated between 4 andthe food sharing with reference to mating behaviors.

5. I used the focal animal sampling method. The target
individuals were one

Table 1. Papaya fruit sharing by Bossou chimpanzees.
Case

Next

The amount of papaya fruits

Sharing

Date

Donor

Recipient

Estrousconsortship

A

B

C

D

E

type

1-1

1 Oct 2002

FF

Pm

No

Yes (estimated)

2

1/2

11/2

1/4

―

Active1-2

1 Oct 2002

FF

Pm

No

Yes (estimated)

―

3/4

1

1/4

11/2

ActiveNo.

2-1

26 Oct 2002

FF

Pm

No

Yes

2

1/3

12/3

1/3

―

Active2-2

26 Oct 2002

FF

Nn

Yes

―

―

11/6

1/2

1/4

15/12

Recovery3

8 Dec 2002

FF

Pm

No

Yes

2

α

2-α

1

1

Passive4

10 Dec 2002

FF

Pm

No

Yes

2

α

2-α

1

2*

Passive516 Dec 2002

FF

Pm

No

Yes

2

α

2-α

1

1

Recovery6

13 Jul 2004

YL

TA

―

―

2

α

2-α

1

1

Recovery7

21 Sep 2004

FF

Pm

Yes

―

2

α

2-α

1-α

1

Passive

Note: FF was alpha male in 2002, and beta male in 2004. YL was alpha male, and TA was gamma male in 2004. Pm andNn were cycling females during my study period. When the recipient was a cycling female, her estrous condition wasnoted. Moreover, if the female who received foods from a male donor was not estrous, I checked whether the donor
engage in a consortship with the female on the next estrous periods. Case 1-2 and 2-2 occurred after other cases ofsharing behavior (Case 1-1 and 2-1, respectively). The amount of fruits was measured by visual estimation: A = Theamount of fruits the donor obtained before sharing, B = The amount of fruits the donor ate before sharing (α means a
few bite), C = the amount of fruits just before sharing, D = the amount of shared fruits, E = the total amount of fruits
that the donor consumed. ― = not applicable. I categorized sharing type by using definition of a previous study3. “Active”
means that “the donor gave part of its food to the recipient after cutting off a piece”. “Passive” means that “the recipient
took part of the food that A was holding, and A made no movement to facilitate the action”. “Recovery” means that “therecipient took part of the food that the donor had placed on the ground”. * After sharing, FF got one more fruit in thevillage and consumed by himself.

alpha male in Period
1, and three adultmales in Period 2.
When chimpanzees
got papaya fruits, I
recorded the timingof food sharing andthe amount of fruits
by visual estimation.
Sharing

attitudes(active/passive)

werecategorizedunder the definitionof a previous study3.

15

Pan Africa News, 14(1), June, 2007

were not estrous. However, the donor seemed tosuccessfully take the females to a peripheral area asconsortship during their next estrous period. I confirmed
the consortship by direct observation in 4 cases (Case 2-1,
3, 4, 5), and for the other 2 cases I estimated consortshipfrom daily attendance record (Case 1-1, 1-2).
I reported some cases of papaya fruit sharing,
however, the chimpanzees did not always share fruits withother individuals. For example, on December 1st, 2002, FF(adult male) started to eat a papaya fruit at 15:36. At 15:43,
Ka (adult female) approached FF and stared into his hand.
Fig. 2. Pm (an adult female) took a papaya fruit that FF (anadult male) had obtained (Case 7).

At 15:45, FF moved about 10m and Ka uttered a screamfor about 2 minutes. At 15:49, Ka approached FF but hemoved away. At 15:52, Ka approached again but FF

RESULTS
Seven episodes of papaya fruit sharing were observedin 222 observation days (Table 1, Fig. 2). The donor

moved about 5m. Ka uttered a scream. For a minute, FF
continued to eat the fruit, then uttered a pant hoot andshowed a display behavior.

always got 2 papaya fruits in the village before sharing. In
two out of the 7 episodes, the possessor shared the fruits

DISCUSSION

twice in one episode. It means that sharing behavior

Bossou is unique, with a chimpanzee habitat

occurred 9 times in total. This article reports these 9 cases.

surrounded by human settlements. The crops must be very

Sharing behavior often occurred just after entering

attractive for chimpanzees especially in seasons when foodthe bush from the village. Especially in five of the 9 cases,

is scarce. However, there seemed to be a sex difference inchimpanzees ate only a few bites before sharing. In Case 4,

their attitudes toward humans. All male chimpanzees werea chimpanzee gave one papaya fruit to another individual,

ready to enter the village and they did not have to beg

and he entered the village again to get another papaya.

others to share fruit. This means that sharing among males

In all of the 9 cases, the recipient approached theseldom occurred, unlike the cases of meat sharing in otherpossessor first, and then showed begging behaviors such as

study sites. On the other hand, some female chimpanzees

reaching their arms and uttering a slight grunt voice. I did

were too shy to have access to the village easily. Such a

not confirm any grimace by the possessor. Three of thedifference in accessibility among individuals may be theobserved cases involved “active sharing” of fruit3, where

cause of fruit sharing behaviors at Bossou.

the donor divided it into two (Case 1-1, 1-2, 2-1). Three

Nishida and Hosaka reported that chimpanzees tend

other cases were “passive sharing”. Recipients took a

to pass worse or smaller food when they share food19. In

portion of the fruits that donors were holding, and the

my observation of 9 cases, chimpanzees always shared lessdonors made no movement either to facilitate or to avoid

than equal half of the fruits. However, they shared fruits inthe action (Case 3, 4, 7). The remaining three cases were

early phase before feeling full. This means that they gave

examples of “recovery”. Recipients took a portion of fruits

up fruits still of value to them. Why does food sharingthat the donor placed nearby (Case 2-2, 5, 6).

occur? Do donors just want to avoid begging or

In all cases except one, donors shared the fruits withcycling females. In two cases, the cycling female washarassment17, 22? Maybe not. Begging is not always
rewarded (e.g., the FF-Ka episode mentioned above).

estrous, and I confirmed mating behaviors during thisThen why did the Bossou males share such valuable

estrous period. In the other six cases, the cycling females

fruits with non-kin others? Here I postulate that papaya

16
sharing contributed to an increase in his chances of
copulations in the future. First, eight out of 9 cases werefrom adult male to cycling female. Especially Pm (adultfemale) was often observed to receive papaya fruit from FF.
In 6 cases, Pm was not estrous at the time. However, FF
successfully took her to a peripheral area for the next
estrous period. In the Bossou group, there were 4-5 cycling
adult females during the study period. However, all but 2
females had not given birth for more than 8 years.
Moreover, one of the 2 females is the mother of FF. The
number of sexually receptive females was so limited at
Bossou. FF may have tried to make a good relationshipwith Pm, even if she was not estrous.

Pan Africa News, 14(1), June, 2007
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Acknowledgments. I thank Professor T. Matsuzawa for his
supervision and advice. Thanks are due to IREB, colleagues of
KUPRI-International team and the local assistants. This study was
financed by a grant from MEXT 16002001 to T. Matsuzawa, agrant under Research Fellowships of the JSPS for Young
Scientists (#160896) and a grant of the Kyoto University
Foundation to G. Ohashi.

18. Matsuzawa T 2006. Sociocognitive development inchimpanzees: a synthesis of laboratory work and fieldwork. In:
Cognitive Development in Chimpanzees, Matsuzawa T,
Tomonaga M, Tanaka M (eds), Tokyo: Springer, pp. 3-33.

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