Pan Africa NewsThe Newsletter of the Committee for theCare and Conservation of Chimpanzees, andthe Mahale Wildlife Conservation SocietyVol. 11 , No. 2

DECEMBER, 2004
P. A. N. EDITORIAL STAFF

Contents:

Chief Editor:
Toshisada Nishida, Japan Monkey Centre, Japan<NEWS>"Kyoto Declaration on Great Apes" has been

Associate Editors:

Adopted.

Christophe Boesch, Max-Planck Institute, Germany

T. Nishida

Jane Goodall, Jane Goodall Institute, USA
Takayoshi Kano, Kyoto Sangyo University, Japan
Tetsuro Matsuzawa, Kyoto University, Japan<NOTES>

William C. McGrew, Miami University, USA

Report on the Chimpanzees of the Wansisi andJohn C Mitani, University of Michigan

Makomayo Areas, Tanzania.

Vernon Reynolds, Oxford University, UK

H. Ogawa, J. Moore, M. Kanamori & S. Kamenya

Yukimaru Sugiyama, Tokai-Gakuen University, Japan
Shigeo Uehara, Kyoto University, Japan

2

3<NOTES>

Richard W. Wrangham, Harvard University, USA

Genito-genital Rubbing among the
Chimpanzees of Mahale and Bossou.

Editorial Secretaries:

K. Zamma & S. Fujita

Noriko Itoh, Kyoto University, Japan

5

Koichiro Zamma, Kyoto University, Japan
Michio Nakamura, Kyoto University, Japan<NOTES>
Predation on a Monkey by Savanna

Instructions to Authors:
Pan Africa News publishes articles, notes, reviews, forums,

Chimpanzees at Fongoli, Senegal.
M. Gašperšič & J.D. Pruetz

news, essays, book reviews, letters to editor, and classified

8

ads on any aspect of conservation and research regarding
chimpanzees (Pan troglodytes) and bilias (P. paniscus). Contributors are requested to write in English and the papers
except forums, reviews and essays should usually be 1,000
words or less. Manuscripts should be submitted by e-mail to:

pan_ed@jinrui.zool.kyoto-u.ac.jp

Photos and figures, however, should be sent by air mail to:T. Nishida, Japan Monkey Centre, 26 Kanrin, Inuyama, Japan.

line for manuscripts is the end of April.
Printed on recycled paper

Increased Hunting of Yellow Baboons (Papio
cynocephalus) by M Group Chimpanzees at
Mahale.

NEW!

Publication of the next issue will be June 2005.

<NOTES>Dead-

H. Nishie

10

Pan Africa News, 11(2), December, 2004

2
<NEWS>“Kyoto Declaration on Great Apes” hasbeen Adopted
Toshisada Nishida
Japan Monkey Centre

The international symposium titled “African
Great Apes: Evolution, Diversity andConservation” was held in Kyoto from 3 – 5
March 2004. This international conference
forms a part of the 21st Century COE Program(A14) “Formation of a Strategic Base for theMultidisciplinary Study of Biodiversity” fundedby the Ministry of Education, Culture, Sports,
Science and Technology. The symposium wasorganized by myself with Prof. Juichi Yamagiwa
as the Secretary-General and Dr. Shigeru
Suzuki as the Leader of the Task Force. The
symposium was opened at 13:00 pm, startingwith poster presentation and welcome party at
the Kyoto Garden Palace Hotel where invited
speakers were staying. For the following
two days, the venue was moved to KyodaiKaikan where oral presentations were given
by 21 invited speakers. It was divided into
six sessions; (1) Behavioral Diversity andOrigins of Human Culture, (2) Sympatry andCoexistence of African Great Apes, (3) Natural

Environments and Evolution of African Great
Apes, (4) The Significance of Pan Social
Behavior, (5) Contributions of African Great
Apes to Biodiversity, (6) Great Apes as the World
Heritage Species. Please check the homepageof the 21st Century COE program (http://ethol.
zool.kyoto-u.ac.jp/coe/coe.htm) for detail.
I reported the establishment of GRASP-Japanin which Japanese researchers of the great apes
cooperate in saving them from the brink ofextinction. The Kyoto declaration on great apes
was also adopted. Several members including
Dr. John Oats, Dr. Richard Wrangham and myself
prepared the draft (see [Column 1]). Sixtynine participants including all field researchersof great apes present signed for this declaration.
Names of signatories are listed in the [Column 2].

Pan Africa News, 11(2), December, 2004
[Column 1] Kyoto Declaration on
Great Apes (Kyoto, March 5, 2004)
As a participant in this symposium, I agreewith the following statement to promoteconservation efforts for the great apes.
1. The international scientists assembled
in Kyoto agreed that many populationsof great apes (humans’ closest living
relatives) are now very seriously
endangered with extinction and that
their numbers continue to decline.
2. As scientists studying great apes
(Chimpanzees, Bonobos, Gorillas
and Orangutans), we feel a special
responsibility to address this issue.
3. We urge that the people and governments
of ape-range and non-range countries work
together to improve the conservation of
the great apes and their natural habitats.
[Column 2]: List of signatories( i n a l p h a b e t i c a l o r d e r )Kaoru ADACHI, Yoko AKARI, Karl AMMAN,
Dean P ANDERSON, Tamotsu AOKI, A Basabose
KANYUNYI, Juliane BRAUER, Takisato DO,
Sarah DUNPHY-CELLI, Tomoo ENOMOTO, Shiho
FUJITA, Fumio FUKUDA, Takeo FUNABIKI,
Takeshi FURUICHI, Miya HAMAI, Chie Hashimoto,
Shuhei HAYAISHI, Kazuhiko HOSAKA, Michael
A HUFFMAN, Kevin D HUNT, Gen-ichi IDANI,
Yumi IKEDA, Agumi INABA, Noriko ITOH,
Mitsuo IWAMOTO, Naomi KAWAGUCHI, Yuichiro KITAYAMA, Mieko KIYONO, Nobuyuki
KUTSUKAKE, Takeshi KOBAYASHI, Suehisa
K U R O D A , Ta m a k i M A R U H A S H I , M i k i
MATSUBARA, Akiko MATSUMOTO, William
C McGREW, John MITANI, Akio MORI, Miho
NAKAMURA, Masato NAKATSUKASA, Toshisada
NISHIDA, Tuyoshi NISHIMURA, John F OATES,
Gaku OOHASHI, Andrew J PLUMPTRE, Melissa J
REMIS, Naruya SAITO, Masaki SHIMADA, Yukiko
SHIMOOKA, David SPRAGUE, Craig STANFORD,
Kazusa SUGAYA, Yukimaru SUGIYAMA, Akira
SUZUKI, Tetsuya SUZUKI, Yukio TAKAHATA,
Yuji TAKENOSHITA, Chihiro TANAKA, Ichiro
TANAKA, Yasuko TASHIRO, Sayaka TUTUMI, Fransde WAAL, Mikihiko WADA, James V WAKIBARA,
Roman WITTIG, Richard WRANGHAM,
J u i c h i YA M A G I WA , S h i n - y a YA M A M O TO ,Shin-ichi YOSHIHIRO, Koichiro ZAMMA

3<NOTES>
Report on the chimpanzees of the
Wansisi and Makomayo areas,
Tanzania
Hideshi Ogawa 1, Jim Moore 2, Masaomi
Kanamori, and Shadrack Kamenya 3
1

Chukyo University, 2University of California, San Diego,
3

Gombe Stream Research Centre

After Kano’s survey on the distribution ofchimpanzees (Pan troglodytes) in Tanzania
between 1965 and 1967 1 , the chimpanzee
population in the Wansisi area has not beenstudied and their current status is unknown.
Ogawa and Kanamori made a brief survey in the
Wansisi area in Aug. 2001. Moore and Kamenya
visited the Makomayo area in July 2001.
Study site and period
The Wansisi area is located approximately 30km

4west of Mpanda and 80km east of Mahale (Fig.1).
This area is approximately 500km2 surrounded by
Kapanga village on the north (6°19'S), Maganje(Mgange) Mts. on the west (30°27'E), Wansisi
Mts. on the south (6°38'S), and Lugala Mts. on
the east (30°43'E)1. The majority of this area is"miombo" woodlands dominated by Brachystegia
and Julbernardia, with bamboo thickets andcultivated fields in the plains and grasslands
along the Kalulu River (1,200m elevation).
On Aug. 3, 2001, Ogawa and Kanamori left
Mpanda, passed Katuma village, turned left at
Kapanga (6°19'S, 30°37'E) out of Mpanda-Mwese
road, and visited Kasunga village (6°21'S, 30°29'E). We walked a total of 30.8km (rounded-trip)at Mgange Hill (6°23' S, 30°32'E) (1,753km) and
Kalole Hill (6°22'S, 30°30'E) on Aug. 4 and 5.
" M a k o m a y o " a s u s e d h e r e r e f e r s to theregion near a series of peaks (Makoma, Sitebi
Makomayo, Mwelu and Kalumangile) locatedabout 25km NNE of Katuma village. All exceed
1,950m elevation (above the tree line) (Fig.1).
The slope northward is gentle, but the mountains
fall steeply in other directions where they are
deeply cut by valleys containing permanent
water and riverine forest; ridges are covered withshort grasses, herbs, some Protea and at exposed
locations, bare rocky earth. This network of highaltitude riverine forest is spread over an area ofroughly 100km2. Below tree line, the surrounding
forest is miombo woodland broadly similar to thatof Wansisi, with large stands of Uapaca in places.
Moore and Kamenya camped at the Lufilisi
River about 12km north of Sitebi Makomayo
Hill from July 23-26, searching for chimpanzees
near 6°07'S, 30°39'E and 5°58'S, 30°42'E.
Results
1. Wansisi:
We did not encounter chimpanzees, but
found 78 beds and 17 feces of chimpanzees
on the slopes and the ridges in the woodlando f M g a n g e a n d K a l o l e H i l l s . We f o u n d
few traces of chimpanzees in the bamboo
thickets and the woodland in the plains.

Pan Africa News, 11(2), December, 2004
Feces of chimpanzees contained seeds ofFicus sp., Grewia mollis, Azanza garkeana,
Pterocarpus tinctorius, and Pseudospondias
microcarpa, leaves of Monocotyledoneae,
and fragments of adult ants (Formicidae).
We also found 3 wadges of bamboo shoots.
Bed groups, in which nests of the same age
were located in the same place, averaged 5.2
beds (range: 2-9, n=15). Fresh nest leaves
indicated that two recent sleeping parties werecomposed of 5 and 8 individuals. Although
there were few evergreen forests on the hills,
some deciduous trees had leaves in this period.
Chimpanzees made beds on such deciduous treesof Brachystegia bussei, B. boehmii, Julbernardia
paniculata, and Isoberlinia angolensis. We saw
claw-marks of a carnivore like a leopard (Panthera
pardus) on the trunk of a nest tree, and diarrhea
feces of the chimpanzee were under that bed.
The population density was roughly estimatedat 0.06 individuals/km 2 based on 20 beds
within 70m of each side of our 19.2km walking
route on the hills, assuming a bed disappears
in 180 days and 70% of beds in the range arefound (after Kano1). This indicates that about
30 chimpanzees inhabit the 500km 2 area.
During the survey, we heard vocalizations ofyellow baboons (Papio cynocephalus) and a
bushbuck (Tragelaphus scriptus), and saw tracesfrom elephant (Loxodonta africana), zebra (Equus
burchellii), and bushpig (Potamochoerus porcus).
2. Makomayo:
No evidence of chimpanzees was encountered.
Although there were a few farms visible in thevalley bottoms, they were not easily reachable
and no local interviews were conducted. The
topography was excellent for hearing sounds, andthe combination of no bed sightings or chimpanzee
vocalizations is a strong indication that none
were present in the high part of the hills at least.
Although there were large areas of evergreen
forest, there were few fruit and no ﬂowering Saba
was visible from lookout points above the valleys.
A number of other primates were encountered.

Pan Africa News, 11(2), December, 2004
Most notably, there were red colobus (Colobus
badius) even in a small (c. 1km 2 ) evergreen
pocket at about 1,900m (to reach other forest
would require crossing at least 500m of shortgrass and exposed rocky soil). This group had
at least 2 adult males; and over 25 individuals
were observed, none with ventral infants. Yellow
baboons and blue monkeys (Cercopithecus
mitis) were seen and heard daily. No galagos(Galago spp.) called at night, and one glimpsedmonkey at lower altitude may have been aredtail (Cercopithecus ascanius). There wereindirect signs of elephant (possibly years old),
zebra, bushpig, hyena (Crocuta crocuta),
leopard, and civet (Viverra civetta), andseveral duiker (unidentified) were glimpsed.
Discussion
In the 1960's, Kano 1 estimated that 40-80
chimpanzees over 3-4 years of age in 1 or 2 unitgroup(s) inhabited the 500km 2 of the Wansisi
area. Our survey showed that chimpanzees
still inhabit the northwestern part of this area.
Local Bende people said that chimpanzees arealso present in the eastern part. However, thepopulation density in this area was low: 0.12
individuals/km2 estimated by Kano1 and 0.06 in
this study. In addition, the chimpanzee populationin the Wansisi area seems isolated from otherpopulations1 in Mahale National Park, Kakungu,
Karobwa, and Ntakata2. These findings indicatethat a limited number of chimpanzees inhabit theisolated Wansisi area. We heard that poachers
killed at least one chimpanzee in this area in June
2003. The population of chimpanzees in the
Wansisi area may face extinction by poaching
and commercial logging. Kano did not visit the
Makomayo area because the nearest villagers
stated there were no chimpanzees there, and thisvisit was entirely consistent with that report.
This is mildly surprising, given the surrounding
miombo and extensive evergreen forest. The
most likely explanation is that the compositionof these high altitude forests is deficient inchimpanzee food plants (or, that it is simply too

5cold -- morning temperatures were around 10°C).
Acknowledgements
Ogawa and Kanamori thank T. Kano, G.
Idani, H. Y. Kayumbo, J. Itani, T. Nishida,
T. Nemoto, E. T. Massawe, and their local
assistants for guidance. This study was permitted
by COSTECH and TAWIRI and financially
supported by a Grand-in-Aid for Scientific
Research of MEXT (#09041160). Moore andKamenya thank Emmanuel Mtiti, Frank Mbago,
Yahya Abeid and Steve Smith, and gratefully
acknowledge the logistical assistance ofCharles Stafford and Anglo American Mining,as well as permission from COSTECH. Theirsurvey was supported by National Geographic
Television; thanks to David Hamlin and the crew.
References

1. Kano T 1972. Distribution and adaptation of thechimpanzee on the eastern shore of lake Tanganyika. Kyoto University African Studies 7:37-129
2. Zamma K, Inoue E, Mwami M, Haluna B, Athumani
S, Huseni S 2004. On the chimpanzees of Kakungu,
Karobwa and Ntakata. Pan Africa News 11: 8-10.

<NOTES>
Genito-genital rubbing among thechimpanzees of Mahale and Bossou
Koichiro Zamma1, Shiho Fujita2
1

Primate Research Institute, Kyoto University

2

Faculty of Applied Biological Sciences, Gifu University

Introduction
Genito-genital (GG) rubbing, which is known
as one of the unique behaviors of bilias (bonobo:
Pan paniscus), has the following characteristics:
two females assume a ventro-vental or ventrodorsal position and the younger female thrusts
her pelvis (Yerkes Regional Primate ResearchCenter, Atlanta 1 ), or two females embrace
ventro-ventrally and rub each other’s sexual
skin together (Wanba, Republic of Zaire2, 3). In
captivity, chimpanzees (Pan troglodytes) were6

also observed to perform GG rubbing, but thevariety of positions they exhibited were different
from those exhibited by bilias 4. Among wild
chimpanzees, we observed GG rubbing at theMahale Mountains National Park, Tanzania
and Bossou, Guinea. Here, we present the firstdescription of GG rubbing by wild chimpanzees.
Methods
Behavioral observations were conducted from
August to November 2003 at Mahale by KZ andfrom December 2001 to March 2002 at Bossou by
SF. KZ followed one of nine focal chimpanzees(P. t. schweinfurthii) on each day and recorded
their body contacts with other chimpanzees. SF
followed one of four focal females (P. t. verus) oneach day and recorded their social interactions.
Observations
Case 1: GG rubbing of Mahale chimpanzees
On September 14, 2003, KZ followed a 7-yearold female named Athena. Her small sexual
skin had not shown swelling. At 11:09:38, when

Pan Africa News, 11(2), December, 2004

Athena was resting on the ground, Ruby, an adultfemale with swollen sexual skin, approachedand lay laterally behind Athena. Then Athenamounted Ruby and shook her hip against Ruby’s sexual skin (but Athena’s sexual skin did nottouch Ruby’s) and groomed Ruby. At 11:10:23,
after Athena shook her hip again, she slightly
stood up and rubbed her sexual skin against Ruby’

Fig. 2. GG rubbing positions of Bossou chimpanzees. DD-1: dorso-dorsal position in which females
stand opposite each other with four limbs on the
ground; DD-2: dorso-dorsal position in which a
female stands on two legs, placing hands on a tree
trunk for support; VD: ventro-dorsal position.

Pan Africa News, 11(2), December, 2004

Fig. 1. GG rubbing positions of Mahale chimpanzees. (a) Position 1: Athena is sitting andRuby is lying laterally. (b) Position 2: Athenais lying on her back and Ruby is lying laterally. Athena moves her hips but Ruby does not.

s on a front-back axis (Position 1, Fig. 1a). At
11:10:51, Athena lay on her back and rubbed
her sexual skin against Ruby’s (Position 2, Fig.
1b). Ruby did not shake her hip, but detached itslightly from Athena’s. Athena also inspected thesexual skin of Ruby by gazing at it and licking it.
KZ observed Position 1 once and Position 2 twice.
Case 2: GG rubbing of Bossou chimpanzees
SF observed 12 successful and 5 unsuccessful
cases of GG rubbing during 556 h of observation(Table 1). The successful cases were classified
into three types of positions (Fig. 2). They
occurred in 3 pairs (Vel & Yo, Vel & Nina, and
Vel & Vuavua). All females were adult (Vel, Yo,
Nina and Vuavua were 42, 40, 47 and 10 years
old, respectively) and had experienced parturition.
Females engaged in GG rubbing in the dorsodorsal position in 9 cases and in the ventro-dorsal
position in 3 cases. In the unsuccessful cases, afemale attempted to perform GG rubbing but theother female did not respond to the solicitation.

7
Discussion
The frequency of GG rubbing among wild
chimpanzees, three bouts during a four-monthobservation at Mahale and 12 bouts during
556 h of observation at Bossou, is lower
than that among captive animals, e.g. 310
bouts during 71 h of observation reported byAnestis4. GG rubbing among the chimpanzeesof Mahale was observed only once by T. Nishida(personal communication), but had not been
described because it was not as established
as among bilias or captive chimpanzees.
In spite of the infrequency of this behavior,
five positions of GG rubbing were observed.
Among bilias, the ventro-vental position wasreported as a typical one 1, 2 , while among
captive chimpanzees, the ventro-vental, ventrodorsal and dorso-dorsal positions, as well as theventro-dorsal-like position where the mountee
lay on her side, were reported 4. Among wild
chimpanzees, we did not observe the ventrovental position but did observe the dorso-dorsal
position, in which a female stood in a bipedal
position with the support of a tree trunk (DD-2).
Five hypotheses on the function of GG rubbing
have been proposed: reconciliation, mate
attraction, tension regulation, expression ofsocial status, and social bonding 5. The small
amount of data from our observations precludes
the possibility of testing these hypotheses
exhaustively, but our results do suggest that
GG rubbing did not occur to release tension. In
our observations, GG rubbing occurred whenchimpanzees were resting (3/3 in Case 1 and2/12 in Case 2) and moving (9/12 in Case 2) but
rarely occurred during feeding (1/12 in Case2). In addition, no conflict was observed among
the chimpanzees engaged in this behavior.
It is still unclear whether GG rubbing provides
a mechanism to attract mates. When Vel andNina engaged in GG rubbing (No. 15 in thetable), the alpha male of the group disturbed
their GG rubbing by shaking a branch at a
distance, which, however, did not result inPan Africa News, 11(2), December ,2004

8
heterosexual copulation. Because Nina kept
trying to engage in GG rubbing with Vel (No.
16 and 17) after the male’s intervention, Nina
appeared to have more interest in GG rubbing
with Vel than in copulation with the male.
Anestis4 indicated that GG rubbing could occur
with higher probability as the degree of bonding
between female chimpanzees was higher. In
Case 2, GG rubbing occurred in particular pairs,
including a mother and her daughter, among
9 adult females of the group; these pairs hadhigh affinity in the party composition 6. The
data supported the hypothesis that GG rubbing
reinforces or at least reflects social bonds.
Interest in another female’s swollen sexual
skin might cause GG rubbing among wild
chimpanzees. In Case 1, Athena gazed and licked
the swollen sexual skin of Ruby. Another time,
Athena was also observed to investigate her own
sexual skin. Athena might perform GG-rubbing
to simultaneously fulfill both her interest in theother’s swollen sexual skin and her interest in
her own immature sexual skin. In Case 2, Vel
was involved in all interactions and reachedmaximum swelling when she GG-rubbed. The
initiator of GG rubbing might be interested in
the swollen sexual skin of the recipient (Vel) andthus act upon her interest by rubbing her own
sexual skin against that of the recipient (Vel).
This study was supported by the Basic Research Fund
( # 111111 t o T. N i s h i d a ) a n d t h e G r a n t f o r t h e
Biodiversity Research of the 21st Century COE (A14).

References
1. Savage-Rumbaugh ES, Wilkerson BJ 1978. Sociosexual behavior in Pan paniscus and Pan troglodytes: a comparative study. J Hum Evol 7: 327-344.
2. Kano T 1980. Social behavior of wild pygmy chimpanzees (Pan paniscus) of Wamba:
A preliminary report. J Hum Evol 9: 243–260.
3. Kuroda S 1980. Social behavior of the pygmy chimpanzees. Primates 21: 181–197.
4. Anestis SF 2004. Female genito-genital rubbing in a group of captive chimpanzees. Int J Primatol 25: 477-488.
5. Hohmann G, Fruth B 2000. Use andfunction of genital contacts among female bonobos. Anim Behav 60: 107-120.
6. Sakura O 1994. Factors affecting party size andcomposition of chimpanzees (Pan troglodytes verus)
at Bossou, Guinea. Int J Primatol 15: 167-263.

<NOTES>
Predation on a monkey by savanna
chimpanzees at Fongoli, Senegal
M. Gašperšič and J.D. Pruetz
Institutum Studiorum Humanitatis (ISH), Ljubljana, Slovenija and Iowa State University, Department of
Anthropology, Ames, Iowa

Although forest-living chimpanzees commonly
include vertebrate prey such as monkeys intheir diet 1 , savanna chimpanzees have beenreported only to eat prosimians2. New evidence
from the recently-established Fongoli studysite in Senegal suggests that chimpanzees there
hunt green monkeys (Cercopithecus aethiops
sabeus). The Fongoli community ranges overan area of at least 63 km 2 and based on fecal
analyses, consumes a species of bushbaby
found here probably Galago senegalensis.
In the third year of study of the Fongoli
community, researchers recorded the following:
On 8 June, 2004, MG and a Senegalese fieldassistant, Mbouli Camara, searched an area on
the left bank of the Fongoli stream, a tributaryof the Gambia River in southeastern Senegal, forsigns of chimpanzees. At 7h13 we heard panthoots about 400 m away and about 1 km E fromthe village of Fongoli. We followed the calls
toward the Gambia River, which at that time was
the only available water source apart from a small
spring near the village of Djendji, several km
away. At 7h23 we found a party of chimpanzees
along the empty riverbed of the smaller Fa-Fa
tributary. The party was moving away, about
30-35 m from us, on the ground in bamboo
woodland. Researchers identified ‘Mamadou’,a young adult male of the community, a second
adult male, a female with an infant, and 3 otherPan Africa News, 11(2), December, 2004
immature chimpanzees. We saw 7 chimpanzees
on the ground, but heard more pant-hoots fromthe surrounding dense vegetation. The three
immature individuals and both males vocalized
and moved in circles around an object on the
ground, which was not recognizable from a
distance. The observed behavior involved
agonistic chasing, as interpreted by JDP after
viewing video footage of the event. After 3
minutes the chimpanzees noticed the observers
and retreated to the nearby woodland, emitting
soft pant-hoots. We waited for 10 minutes andthen approached the site along the dry streambed.
There we found a dead green monkey, apparently
a young female. The skull was completely
fractured in the frontal, parietal, and occipitalregions, exhibiting a radiating fracture patternindicative of blunt force trauma. The patternof fracture suggests that the monkey was killedby hitting its head against a hard substrate, as
has been observed at Tai Forest3. The brain waspartly missing, and the blood was fresh and notyet dried. The monkey had wounds on the neckand shoulders; the left leg had no ﬂesh left on it;
both right limbs were missing, apparently having
been pulled off. The abdominal area was ripped
open, with intestines spilling out. The victim
appeared freshly killed, as there eas no odor of
decay; there was no maggot activity, and few ﬂies
had arrived at the corpse. We waited nearby for
30 minutes, in case the chimpanzee party cameback, but they did not. We returned to the spot
the next day to examine the body, but it was gone.
MG filmed the encounter, but the actual huntor kill and the monkey cannot be seen due
to the distance and dense vegetation. Given
the evidence, both behavioral and physical,
we conclude that this is the first evidence ofmonkey hunting by savanna chimpanzees ofwest Africa. Primate prey species commonly
hunted by chimpanzees at other sites, such ascolobus monkeys, do not occur in this semiarid environment (Guinean and Sudo-Guinean
vegetation types). We suspect that vertebratehunting by Fongoli chimpanzees is opportunistic

9
and probably not cooperative, as the preyspecies is the most terrestrial African guenon.
Other potential chimpanzee prey species that
occur at Fongoli include the patas monkey(Erythrocebus patas) and Guinea baboons (Papio
papio), as well as smaller ungulates such as
bush duiker (Cephalophus sp.), oribi (Ourebia
ourebi), warthog (Phacochoerus aethiopicus),
and bushbuck (Tragelaphus scriptus). Encounters
between patas monkeys and chimpanzees have
not yet been observed. Agonistic encounters
between chimpanzees and baboons have been
observed during the peak of the dry season whenboth species are drawn to a few available water
sources in the study area. Contest competition
between chimpanzees and baboons over water
has been recorded, with chimpanzees acting as
aggressor but baboons retaliating in large groups;
baboons acting as aggressor, with chimpanzees
retreating; and, at other times, chimpanzees
and baboons drink sequentially at the water
source, with no overt competition. Baboons
have also been observed to avoid the approachof chimpanzees as signalled by pant-hooting.
With increased habituation of the
F o n g o l i c o m m u n i t y, w e h o p e t o c l a r i f y
the hunting behavior of chimpanzees insuch an open habitat as well as patterns ofmeat sharing and other related behaviors.
We acknowledge funding by the U.S. Fish
and Wildlife Great Ape Conservation fund,permission to work in Senegal grantedby the Eaux et Forets du Senegal and theRepublique du Senegal, and field assistanceby Mbouli Camara and Dondo Kante.
References
1. Stanford C 1998. Chimpanzee and Red Colob u s : T h e E c o l o g y o f P re d a t o r a n d P re y .
Harvard University Press, Cambridge.
2. McGrew WC., Baldwin PJ, and Tutin CEG.
1988. Diet of wild chimpanzees (Pan troglodytes verus) at Mt. Assirik, Senegal. I. Composition. Amer J Primatol 16: 213-226.

Pan Africa News, 11(2), December, 200410
3. Boesch C, Boesch-Achermann H
2 0 0 0 . T h e C h i m p a n z e e s o f t h e Ta ï F o r est. Oxford University Press, Oxford.

<NOTES>
Increased hunting of yellow baboons(Papio cynocephalus) by M groupchimpanzees at Mahale
Hitonaru Nishie
Kyoto University

Introduction
Since the first observation by Nakamura 1, no
one has reported cases of predation on yellow
baboons by the M group chimpanzees at Mahale.
Nishida2, 3 reported that some baboon groups had
invaded the range of the M group chimpanzees
from the early 1990s, and direct encounters
between chimpanzees and baboons have been
observed (N. Itoh, T. Matsusaka, M. Nakamura,
K . Z a m m a & T. N i s h i d a , u n p u b l i s h e d ) .
Furthermore, many authors 3, 4, 5 have reported
the importance of indirect competition
between baboons and chimpanzees. However,
it is still unclear how possible competition
influences the relationship between them.
There have been no reports of predation
on baboons by chimpanzees at other sites,
aside from Gombe where olive baboons(Papio anubis) were formerly preyed uponby chimpanzees 6 . This variation in preyselection among chimpanzee populations
may be explained partly by differences inecological factors and partly by local traditions6.

Observations
Case 1: 2 March 2004 (rainy season)
M group chimpanzees dispersed into small
parties. At 9:35, I found Alofu, the alpha male,in a tree near the Sinsiba tourist camp, holding
the carcass of an infant baboon. Other than
Masudi (adult male) and Jidda (juvenile female),
there were no other chimpanzees in the area. A
tourist guide who had arrived earlier observed
Jidda capturing and releasing the infant baboon
and then Alofu capturing it immediately. Mostof the prey’s body was intact when I arrived.
At 9:38, Alofu, holding the carcass, allowed
Masudi to bite off some meat from it. At 9:45,
Masudi approached again, but this time turned
his back on Masudi. Jidda stayed lower, picking
up some bone pieces and sucking on them. At
9:57, Masudi approached Alofu again, and thistime Alofu let him bite the meat. At 10:01,
Masudi tore a leg of the prey off and moved
away. Then both males continued to eat the meat.
At 10:21, Jidda climbed to the ground andsought out bone pieces. At 10:24, Masudicame down. Masudi had already consumed
a whole leg. Alofu followed Masudi
and sometimes stopped to eat the meat.
At 10:34, Alofu resumed eating the meat,
while Masudi and Jidda moved on. At 10:46,
Alofu left and caught up with Masudi and
Jidda, later stopping to eat the meat. At 10:50,
the three chimpanzees moved into a swamp,
ending my observation. At 10:55, Alofu wasseen no longer carrying the meat. Therefore,
I could not confirm whether Alofu had
consumed the meat entirely or abandoned it.

Pan Africa News, 11(2), December, 2004

Figure

1.

Alofu

11

capturing

Case 2: 24 September 2004 (dry season)
M group chimpanzees ranged in a large party
that included most members. At 8:11, I heard
chimpanzees nearby. At 8:14, my focal female
suddenly turned and headed for their commotion.
At 8:25, Michio (juvenile male) on a tree
near the activity grasped a juvenile yellow
baboon and released it at once, and then Alofu
immediately captured it (Figure 1). Alofu came
down from a tree and dragged the prey on the
ground. Alofu sometimes let the prey run away
and soon recaptured it. The prey, bleedingslightly from the abdomen, was still alive.
There were at least 15 chimpanzees in the area;some followed Alofu, others just stared at him.
At 8:37, Alofu abandoned the prey without
killing it. Then Cadmus (young male) captured it
and climbed a tree. At 8:39, Cadmus swung theprey and hit it against a branch, which appeared
to kill it. At 8:42, Cadmus returned to the groundand slapped and stamped the prey. At 8:44,
Cadmus climbed up a tree. Then Orion (youngmale) came and stared at Cadmus. At 8:46,
Michio climbed the tree and approached Cadmus.

ajuvenile

baboon.

Cadmus began eating the baboon. At 8:50, Orion
moved away. At 8:55, Cadmus swung the preyand dropped it on the ground. Michio approached
to pick it up but was charged by Cadmus. Cadmus
climbed the tree and resumed eating the meat.
At 9:05, Darwin (young male) climbed
up the tree to approach Cadmus. Darwin
began to perform pant-hoot. Then,
Darwin began to whimper and screamnear Cadmus, who was eating the meat.
At 9:44, Cadmus dropped the carcass, andDarwin picked it up. Cadmus climbed down andleft. Only the area around the prey’s head wasconsumed. At 9:50, Christina (adult female)arrived with her daughter and mated with Darwin.
At 9:55, Xtina picked up a bone piece and left.
At 9:59, Darwin dragged the carcass. At 10:07,
Darwin resumed eating the prey’s legs and arms
in a tree. At 10:45, Darwin released the carcass
and began eating fruit. At 10:48, Darwin swung
the carcass and dropped it. Darwin came down
to pick it up. At 10:50, Darwin abandoned
the carcass and moved away. The carcass had
been consumed around the head, part of the12abdomen, some parts of the arms, and both legs.
Discussion
The frequency of chimpanzee hunting ofbaboons has increased recently at Mahale(Table 1). This increase is apparently causedby the increasing number of yellow baboonsin the M group range 1, 2, 3, 4 . However, thiscannot explain the diachronic changes inchimpanzee attitudes toward baboons.
In Case 1, all attendants (Alofu, Masudi,
and Jidda) seemed eager to get the meat of theinfant baboon. Alofu, the holder, kept the prey
for a long time and was reluctant to share themeat with Masudi. This indicates that at least
the chimpanzees of the M group sometimes
have the desire to eat the meat of baboons.
In Case 2, however, the first holder, Alofu,
released the juvenile baboon without killing it.
Moreover, although more than 15 chimpanzees
were present, just a few begged for the meat, andthese did not persist for long. The other holders,
Cadmus and Darwin, also ate just a little of themeat and eventually left much of it uneaten.
What accounts for this difference? It is
likely that the size of the prey influenced theattitude of the chimpanzees. In the case of
hunting red colobus, the main prey of the M
group, chimpanzees also show a preference forsmall prey rather than larger prey. The preysize of Case 2 was slightly larger than thatof Case 1 and thus, perhaps, less attractive.
Moreover, the variation in chimpanzee attitudes
toward baboons might indicate the flexibilityof the behavior, which has not yet become
customary among the group’s members. It is
important to collect variations in behavioral
repertoires among chimpanzee populations tocharacterize chimpanzee cultures; however,gathering detailed descriptions of diachronicchanges in behaviors within a chimpanzee
unit-group is even more important forunderstanding how chimpanzees experience
their own “familiar” and “usual” world.

Pan Africa News, 11(2), December, 2004
This study was supported by the Basic Research Fund
( # 111111 t o T. N i s h i d a ) a n d t h e G r a n t f o r t h e
Biodiversity Research of the 21st Century COE (A14).

References

1. Nakamura M 1997. First observed case ofchimpanzee predation on yellow baboons(Papio cynocephalus) at the Mahale Mountains National Park. Pan Africa News 4(2): 9-11.
2. Nishida T 1997. Baboon invasion into chimpanzee habitat. Pan Africa News 4(2): 11-12.
3. Nishida T 2002. Competition between baboons andchimpanzees at Mahale. Pan Africa News 9: 23-26.
4. Matsumoto-Oda A, Kasagula MB 2000. Preliminary
study of feeding competition between baboons andchimpanzees in the Mahale Mountains National Park,
Tanzania. African Study Monographs 21: 147-157.
5. Uehara S, Ihobe H 1998. Distribution andabundance of diurnal mammals, especiallymonkeys, at Kasoje, Mahale Mountains, Tanzania. Anthropological Science 106: 349-369.
6. Uehara S 1997. Predation on mammals by the chimpanzee (Pan troglodytes). Primates 38: 193-214.
Pan Africa News, Vol. 11, No.2
Published in December, 2004
Address:
Japan Monkey Centre
26 Kanrin
Inuyama, Aichi 484-0081
JAPAN
TEL: (+81)568-61-2327
FAX: (+81)568-62-6823
E-mail: pan_ed@jinrui.zool.kyoto-u.ac.jp

nori