Pan Africa News
Newsletterof theCommittee
Care
TheTheNewsletterof theCommittee
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andandConservationof ofChimpanzees,
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ISSN 1884-751X (print), 1884-7528 (online) mahale.web.infoseek.co.jp/PAN/

DECEMBER 2009
P. A. N. EDITORIAL STAFF
Chief Editor:
Toshisada Nishida, Japan Monkey Centre, Japan
Deputy Chief Editors:
Kazuhiko Hosaka, Kamakura Women’s University, Japan
Michio Nakamura, Kyoto University, Japan
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
Editorial Secretaries:
Noriko Itoh, Kyoto University, Japan
Koichiro Zamma, Hayashibara GARI, Japan
Agumi Inaba, Japan Monkey Centre, Japan

VOL. 16, NO. 2

Contents<NOTE>
Power Takeover Occurred in M Group of the Mahale
Mountains, Tanzania, in 2007
Agumi Inaba

13<NOTE>
Savanna Chimpanzees (Pan troglodytes verus) Prey on Patas
Monkeys (Erythrocebus patas) at Fongoli, Senegal
Jill D. Pruetz & Joshua L. Marshack
15<NOTE>
Aesthete in the Forest? A Female Chimpanzee at Mahale
Collected and Carried Guineafowl Feathers
Michio Nakamura

17<NOTE>The Death of a Newborn Chimpanzee at Mahale: Reactionsof its Mother and Other Individuals to the Body
Takanori Kooriyama
19<FORUM>
Foreword
William C. McGrew

An Encounter between a Leopard and a Group of
Chimpanzees at Gombe National ParkAnn H. Pierce

22
22Instructions for Authors:
Pan Africa News publishes articles, notes, reviews,
forums, news, essays, book reviews, letters toeditor, and classified ads (restricted to non-profit
organizations) on any aspect of conservation andresearch regarding chimpanzees (Pan troglodytes)
and bilias (Pan paniscus). Contributors arerequested to write in English and the papers except
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words or less. Articles and notes will bepeer-reviewed by at least one appropriate expert onrequest of the PAN editorial staff.
Manuscripts should be formatted as DOC or RTF
files and submitted by e-mail to:pan.editor@gmail.com
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GIF files and sent separately by e-mail attachments.
PAN is published twice a year in June andDecember. Deadline for manuscripts is one month
before publication (i.e. the ends of May andNovember).

<NOTE>

Power Takeover Occurred in
M group of the Mahale
Mountains, Tanzania, in 2007
Agumi Inaba
Japan Monkey Centre, Japan

Male chimpanzees compete for high rank, especially,
alpha status since it confers the high reproductive
success1,2. Power takeovers have been recorded, butopportunities to observe such incidents are rare. Social
intelligence seems to be exhibited so much in the courseof dominance competition that documentation of social
interactions is theoretically important3–6. I briefly report
here some interesting episodes during the recent takeover

13

14

Fig. 1. Alofu, the ex-alpha male, in September 2007.

of alpha status by a young adult male.
Since 2003, in the M group of Mahale Mountains
National Park, Alofu (25 years old, Fig. 1) had kept hisalpha status. In 2007, however, the alpha male waschanged. The beta male, Pim (19 years old, Fig. 2),challenged Alofu and then succeeded in seizing the alphastatus. Back in November 2003, Fanana (alpha male, 25
years old at that time) suddenly disappeared, and Alofu(beta male, 21 years old at that time) advanced his status
to alpha7. Alofu thus remained in power for three years
and six months. However, according to Dr. Michio
Nakamura, who stayed at Mahale in June 2007, Pim

Pan Africa News, 16(2), December 2009

Fig. 2. Pim, the current alpha male, in September 2007.

ceased to pant-grunt to Alofu during that time. It seemedthat the dominance relation between them became
ambiguous. Subsequently, they seemed to fight eachother, scrambling for political power up to August, but
the details were unknown between June and August.
I began to study the social relationships among theadult males of M group on August 10 and ended myobservations on December 22, 2007. On August 17, 2007,
Dr. Toshisada Nishida and I observed Alofu pant-grunting
to Pim for the first time, and thus we speculated that Pim
had taken over the alpha status around that time, or notlong ago. Alofu’s display dragged out for a long time. In
contrast, Pim who was young and rough withpiercing eyes, often performed short but
vigorous charging displays. After the changeof political power, other males of the M
group behaved with indecisiveness, playing
up to Alofu but following Pim as well when
he appeared. Alofu was not yet old, and he
was still strong enough to retaliate against
Pim and regain the alpha status. Nevertheless,
Alofu eventually seemed to be satisfied withthe beta male position. In fact, the real threat
to Pim may have come from the younger
rivals who had matured well. Primus (16years old) was (and is) the principal rival,
with a baby face and well-built physique,threatening Pim and Alofu. Cadmus (16years old) is small in size but a good athlete.
We would expect them to be promising
candidates for the top position.
Youngsters were active, but a senior,Kalunde (about 44 years old), continued to
be strong and influential in politicalsituations. On August 28, 2007, males
showed opposition to Pim over a cycling
Fig. 3. Kalunde (center) began to impose himself in the agonistic relationshipfemale. Although Pim solicited support frombetween Alofu and Pim. When Alofu (right) charged toward Pim (left), Pim
Kalunde more than once, Kalunde switchedescaped (August 2007, from video clip by Dr. Takanori Kooriyama).
his allegiance to Alofu as soon as he sawPan Africa News, 16(2), December 2009
Alofu rushing toward Pim, and he also gave chase to Pim(Fig. 3). Pim escaped at full speed, then returned to
Kalunde hastily, and finally hugged Kalunde to make up
with him. On November 13, when Pim was defeated by
the team of Primus-Cadmus, other males were sitting andwatching at a distance, and only Kalunde walked to Pim’sside. These incidents did not have an impact on Pim’s
status; the power balance among highest-ranking males
seemed to depend not only on the attitude of many adultmales, but also on Pim’s assertiveness.
There was one male who came back to the central
part of the group, as if he had foreseen the change inalpha status. He was Fanana (29 years old, Fig. 4), thelast alpha male but one, who had disappeared in 2003 andsince then almost always ranged alone within the M
group’s territory. In August 2007, Fanana repeated somebehavior patterns, such as traveling with members for a
few days and then disappearing for another few days.
Fanana seemed to adapt himself entirely to the group by
September. On November 12, Fanana turned up, after aninterval of several days, at the hill where M groupmembers were resting. Fanana came up quietly and sat in
the middle of the current power coalition. Alofu, Kalunde,
Cadmus and Tarnie (female, estimated 15 years old) had
all been grooming in tandem. Seeing Fanana coming, allfour chimpanzees turned around and sat immediately
surrounding Fanana, and then they began to groom him.
Fanana engaged in vigorous social play, such as
wrestling with other males, as if he were compensating
for previous lost time. Furthermore, he often traveled

15
Basic Scientific Research Fund (#19255008 to T.
Nishida) and the Global Environment Research Fund
(F-061 to T. Nishida) of the Ministry of the Environment,
Japan.

REFERENCES
1. Hasegawa T, Hiraiwa-Hasegawa M 1990. Spermcompetition and mating behavior. In: Nishida T (ed) The
Chimpanzees of the Mahale Mountains. University ofTokyo Press, Tokyo, pp. 115–132.
2. Inoue E, Inoue-Murayama M, Vigilant L, Takenaka O,
Nishida T 2008. Relatedness in wild chimpanzees: theinfluence of paternity, male philopatry, and demographic
factors. Am J Phys Anthropol 137:256–262.
3. de Waal FBM 1982. The Chimpanzee Politics. Jonathan
Cape, London.
4. Nishida T 1983. Alpha status and agonistic alliance in wild
chimpanzees (Pan troglodytes schweinfurthii). Primates
24:318–336.
5. Nishida T, Hosaka K 1996. Coalition strategies among adultmale chimpanzees of the Mahale Mountains, Tanzania. In:
Great Ape Societies, McGrew WC, Marchant LF, Nishida T(eds), Cambridge University Press, Cambridge, pp.
114–134.
6. Hosaka K, Nishida T 2002. Restoration of alpha status tothe ostracized ex-alpha male. In: Mahale Chimpanzees:
Thirty-seven Years of “Panthropology”, Nishida T, Uehara
S, Kawanaka K (eds) Kyoto University Press, pp. 439–471(in Japanese).
7. Nishie H, Nishida T 2005. A new record of reclusion of analpha male chimpanzee and his decline from the alpha status
in the Mahale Mountains National Park, Tanzania. Primate
Res Suppl:S–9 (in Japanese).

<NOTE>

Savanna Chimpanzees (Pantroglodytes verus) Prey on Patas
Monkeys (Erythrocebus patas) at
Fongoli, Senegal
Fig. 4. Fanana coming back with fresh wound on the rightside of his face (September 2007).

with Pim. Although Fanana’s status fell to near the lowest
end of the adult males, it seemed that Pim began to rely
on Fanana to some extent. Based on these initial findings,
I would like to follow the movements of the adult malesof M group.
I thank T. Kooriyama and M. Nakamura forproviding valuable information and video clips; T.
Nishida and K. Hosaka for constructive comments. The
research was supported by the MEXT Grant-in-Aid for

Jill D. Pruetz1 & Joshua L.
Marshack2
1. Departments of Anthropology, Iowa State University, USA
2. Department of Anthropology, Washington University, USA

INTRODUCTION
Chimpanzees across Africa include some meat intheir diet1. In most communities where chimpanzees havebeen studied over the long term, primate prey isapparently preferred over other animal prey, with redcolobus monkeys (Piliocolobus badius) comprising themost common monkey species eaten2. At Fongoli,16
Senegal, it is likely that chimpanzees have never hadaccess to red colobus monkeys at this hot, dry and open
site, although the range of red colobus monkeys (P.b.
temminckii) in Senegal and adjacent countries in West
Africa has diminished3. Fongoli chimpanzees are
currently not sympatric with red colobus, although theyeat vertebrate prey of various types, including a numberof primate species.
We previously reported two of the three primate
species as preyed upon by chimpanzees at Fongoli(Chlorocebus aethiops4, Galago senegalensis5). Fongoli
chimpanzees use modified sticks as jabbing tools to
obtain bushbabies. We have since observed Fongoli
chimpanzees hunting banded mongoose (Mungos
mungo)5 and baboons (Papio hamadryas papio) andfeeding on bushbuck (Tragelaphus scriptus) and as well
(Pruetz, unpublished data). Here, we report on the
inclusion of a fourth primate species eaten bychimpanzees at Fongoli, the patas monkey (Erythrocebus
patas). This is the first recorded incident of predation on
patas monkeys by chimpanzees. Fongoli is the only sitewhere habituated chimpanzees are sympatric with thismonkey species.

STUDY SITE
Chimpanzees in Senegal inhabit the area known as the
Mandingue Plateau, which defines the northern andgeographical limits of chimpanzees’ range in West Africa.
This region of southeastern Senegal is Sudanian savanna
and Guinean woodland. The Fongoli site (12º40’ N, 12º13’
W) is at the junction of the Sudanian and Sudano-Guinean
vegetation belts, which can be envisioned as asavanna-woodland mosaic1. Rainfall averages less than 800
mm annually (Pruetz, unpublished data). In southeastern
Senegal, the rainy season is from June through September.
Other animal species at the site include those associatedwith open environments, such as patas monkeys and oribi(Oribi oribi), as well as species that typically use closed
habitats more extensively, such as green monkeys andbushbucks. Humans that live in the area include the Bedik,
Bassari, Diahanke and Malinke.
The Fongoli chimpanzee community has been studied
since April 2001, and systematic behavioral data collection
began after habituation of adult males in 2005. The
community size ranges between 31–34 individuals annually,
with approximately 10–11 adult males, 7–8 adult females
and varying numbers of immature chimpanzees comprising
the community, based on records collected since allindividuals were identified in January 2006 (Pruetz,
unpublished data). The Fongoli chimpanzee home range isestimated, minimally, to be 63 km² and is predominantly
open woodland and grassland, with small patches of gallery
forest and seasonally cultivated fields1.
OBSERVATIONS
Few encounters have thus far been observed between

Pan Africa News, 16(2), December 2009
patas monkeys and chimpanzees at Fongoli in the four
years following habituation, likely because both
chimpanzees and humans are predators of patas in this area
(Pruetz, personal observation). In the first encounterrecorded between patas monkeys and chimpanzees, during
the 2008 dry season, the latter gave wraa calls at two malepatas (one adult, one subadult) that had been drinking at thepermanent water source near a resting chimpanzee party.
Both patas monkeys fled quickly and one adolescent malechimpanzee gave chase, although he appeared not to run atfull speed. He did not gain on the monkeys at any point, but
the chase ended out of sight of the observer. The monkeys
did not vocalize in any way during this encounter.
In the second encounter observed between
chimpanzees and patas monkeys, in July 2008, two older,adult male chimpanzees attempted to hunt at least two patasmonkeys. The older and lowest ranking of the two males
was successful in capturing a one-year old (juvenile)monkey. The behavior of these two, older adult malechimpanzees suggested a coordinated attempt at patasmonkey predation. During foraging-related travel, one male
stopped immediately after the first, and both males
remained silent and attentive until the same moment when
they raced to the trees in the woodland habitat in which thepatas monkeys were ultimately found. While this may be
explained as coincidental, the silent communication
between the adult males, including the exchange of gaze,
indicated that such behavior was not characteristic of one or
the other’s selfish motivation or desire to reach the monkeys
first. Observers waited approximately 10 seconds before
following the subjects so as to refrain from alerting themonkeys prematurely, so were able to only observe thesecond attempt on a second juvenile patas monkey (one or
two years of age). The older male had already begun eating
the juvenile patas he caught while outside of observers’
view. The other adult male was observed to jump into thetree containing the second juvenile patas monkey and climb
quickly toward it. The patas monkey almost immediately
leapt from the tree (at a height of less than 10 meters) to theground and began running. The adult male chimpanzee
immediately stopped chasing the monkey once it landed on
the ground and returned to the other chimpanzee’s kill. The
monkey was shared extensively between the two adultmales and was consumed entirely save for its bones and tail.
At no time did patas monkeys elicit alarm calls or othervocalizations before or after the predation event.
In the third encounter between chimpanzees and patasmonkeys, in July 2009, observers followed an adult maleand an older adolescent male. The two chimpanzees wereobserved to travel cautiously, apparently listening to sounds
made by patas monkeys. Both chimpanzees gave chase towhat turned out to be a patas monkey, but they were lostfrom view during the hunt and capture. Approximately
eight minutes later, the adolescent male was observed
feeding on a young patas monkey, not sharing with the adultmale until 197 minutes later.

Pan Africa News, 16(2), December 2009DISCUSSION
Fongoli chimpanzees have now been observed tofeed on each of the non-human primates sympatric withthem in their hot, dry and open environment. Patasmonkeys are known to be the fastest terrestrial primate6.
Although the speed of these monkeys was long thought
to be a trait selected for anti-predation in this species, a
recent study in Laikipia, Kenya recorded a variety ofanti-predator strategies in this species, and fleeing frompredators accounted for less than half of all responses7.
As seen in the observed predation attempt by an older
male chimpanzee on a young patas monkey, escape into
the trees did not deter pursuit but escape via running on
the ground appeared to deter pursuit. Notably, monkeys
did not elicit alarm calls in any observed encounters.
Chimpanzees’ ability to pursue their prey in trees
precludes the ability of monkeys to escape safely into
trees as with other terrestrial predators, so that patasmonkeys in the Fongoli region appear to use crypticity as
an initial anti-predation strategy when faced with achimpanzee predator.
ACKNOWLEDGEMENTS
Research at Fongoli is carried out with the kindpermission of the Republic of Senegal and the Department duEaux et Forets and the Arrondissement du Bandafassi. Funding
has been provided by the National Geographic Society, Iowa
State University, Leakey Foundation, Wenner-Gren Foundation
for Anthropological Research, National Science Foundation,
U.S. Fish & Wildlife Great Ape Conservation Grant, Primate
Conservation Inc., and American Society of Primatologists.
Assistance is provided in the field by Dondo Kante, Mboule
Camara, Michel Sahdjaro, and Wally Camara.

REFERENCES
1. Pruetz JD 2006. Feeding ecology of savanna chimpanzees at
Fongoli, Senegal. In: The Feeding Ecology of Great Apes andother Primates, Hohmann G, Robbins M, Boesch C (eds),
Cambridge University Press, Cambridge, pp. 161–182.
2. Mitani JC, Watts DP 2001. Why do chimpanzees hunt andshare meat? Anim Behav 61: 915–924.
3. Galat G, Galat-Luong A, Nizinski G 2009. Increasing dryness
and regression of the geographical range of Temminck’s red
colobus Procolobus badius temminckii: implications for its
conservation. Mammalogy 73: 365–368.
4. Gaspersic M, Pruetz JD 2004. Predation on a monkey bysavanna chimpanzees at Fongoli, Senegal. Pan Afr News 11:
8–10.
5. Pruetz JD, Bertolani P 2007. Savanna chimpanzees (Pantroglodytes verus) hunt with tools. Curr Biol 17: 1-6.
6. Hall KLR 1968. Behaviour and ecology of the wild patasmonkey, Erythrocebus patas, in Uganda. In: Primates:
Studies in Adaptation and Variability, Jay PC (ed), Holt,
Rinehart and Winston, New York, pp. 32–119.
7. Isbell LA, Pruetz JD,Young TP, Lewis M 1998. Locomotory
activity differences between sympatric patas monkeys(Erythrocebus patas) and vervet monkeys (Cercopithecus
aethiops): Implications for the Evolution of Long Hindlimb
Length in Homo. Am J Phys Anthropol 105: 199–207.

17<NOTE>

Aesthete in the Forest?
A Female Chimpanzee at
Mahale Collected and Carried
Guineafowl Feathers
Michio Nakamura
Wildlife Research Center, University of Kyoto, JapanINTRODUCTION
We humans sometimes collect objects that we
consider beautiful and carry them home. For example, we
may collect beautifully colored stones or shells when wewalk on the beach or may pick up beautiful flowers orleaves in the forest to press in a book. We collect suchitems not because they have practical use―we do not eat
them, we do not sell them, nor do we use them as tools or
medicines―but simply to keep them on a shelf.
One item that attracts those who walk in the forestof Mahale, Tanzania, is the beautiful blue-spotted blackfeathers from the crested guineafowl Guttera pucheranithat are often found on the forest floor (Fig. 1).
It is not surprising that chimpanzees (Pantroglodytes) collect and carry things that are directly
beneficial to them, such as food. For example,
chimpanzees at Mahale sometimes collect five or six
lemons from a tree, carrying one in the mouth, one ineach hand and foot, and perhaps one more in the groinpocket. So burdened, they cannot walk very far, but may
just climb out of the tree and eat the fruit nearby. They

Fig. 1. Feathers of crested guineafowls collected by theauthor at Mahale.

18
also carry various food items between their lips, in a handor foot, on the back or head, or in the groin or neckpocket, or drag food with one hand when they have tocatch up with other chimpanzees1. Another example ofcollecting and carrying at Mahale has to do with fishingtools: chimpanzees sometimes prepare these tools atsome distance from an ant nest and carry them for up to
70 meters2.
Immature chimpanzees sometimes carry objects
that are not related to feeding. During play, they oftentransport objects such as twigs1. They may carry twigs,
logs, tree bark, colobus skin, and human artifacts as if
they were carrying “toys” or “dolls” 3–5. Researchersconsider such objects “toys” because juvenile
chimpanzees often handle them in a playful manner, suchas hitting them, tapping them, or putting them on theirhead, while showing a typical play face. Such “toys”
usually do not attract adults, but only infants andjuveniles show interest and often play tug-of-war withthese objects.
Here, I would like to report a case in which an old
female chimpanzee calmly collected several guineafowl
feathers and carried them for more than 30 minutes
without displaying any emotional expressions, such as aplay face.

OBSERVATIONSAt 10:36 am on 11 October 2009, when I wasobserving an adult female called Cynthia, my research
assistant noticed that an old female called Wakusi(estimated to be 48 years old at the time of observation)
was holding two or three guineafowl feathers in her left
hand. As we observed her, she walked a bit with thefeathers in her hand and picked up two more feathers that

Pan Africa News, 16(2), December 2009
happened to be on the trail we were observing. Now
carrying four or five feathers in her hand, she kept
walking as the other chimpanzees headed north. We
followed her to observe the consequences.
At 10:41, Wakusi copulated with a 9-year-old
adolescent male called Emory while holding the feathers
in her lips. Then, we lost sight of her briefly while we
waited for other chimpanzees sitting on the trail to depart.
When we caught up with her at 10:44, she was lying on
the trail with five individuals close by. She still had thefeathers in her left hand, and Emory looked at them.
Cynthia’s 1-year-old infant appeared to become interested
in what Wakusi had in her hand. Wakusi gently put thefeathers in her groin pocket and continued lying on the
trail. The infant went closer to Wakusi to look at her groinarea where she had put the feathers. Wakusi took out thefeathers with her left hand (Fig. 2) and then returned themto her groin pocket. The infant looked at Wakusi’s belly
again and then went to her backside. When the alphamale Pimu began to groom Wakusi, the infant went toWakusi’s ventral side and took the feathers out of her
groin pocket. Although Wakusi moved her hand toward
the baby slightly, she did not attempt to retrieve thefeathers. The infant sniffed and licked the feathers, but
discarded them after a while when its mother was about
to depart.
At 10:51, when the alpha male stopped grooming
and left, Wakusi was left alone for a while. As she got up,
she picked up three feathers one by one with her hand,
put them between her lips, and departed into the bush. As
she moved through the bush, she again presented to
Emory and copulated with him, still holding the feathers
in her lips. As she and other chimpanzees moved on in
the dense bush, we could not observe her continuously,

Fig. 2. Wakusi lying supine with the guineafowl feathers in her hand. Cynthia’s baby is sitting in front of her facing Wakusi. In theline drawing of Wakusi, the feathers are painted black.

Pan Africa News, 16(2), December 2009
but when we caught up with her, she had the feathers in
her hand, between her lips, or in her groin pocket.
At 11:12 am, Wakusi and some others departed insome haste, and again we lost sight of her because otherchimpanzees were barring our way. At this time, she still
had the feathers in her mouth. When we reached the trail
where she had just passed, I found one guineafowl
feather on the ground; its proximal end was wet withsaliva (Fig. 3). When we found Wakusi with a party ofchimpanzees feeding on a Ficus tree at 11:35, she nolonger had the feathers.

19
handled them in a playful manner (so, they could havebeen a “toy” to the baby), Wakusi herself never attempted
to play with them or showed a play face. Consequently, it
is difficult to say that Wakusi considered them toys in theusual sense.
The old female chimpanzee just calmly collected
and carried several bird feathers. It is difficult to guess
whether she found them beautiful, as humans do. She did
not seem to collect the feathers for a particular functional
reason. Consequently, can we say that she collected them
just because she liked them? Many archeologists assume
that art or symbolic representation only appeared inmodern humans about 50,000 years ago7. However,
because the underpinning aesthetic mentality cannot be
fossilized, it is possible that similar aesthetic sensibility
had evolved much earlier in the chimpanzee-humanclade.

REFERENCES

Fig. 3. A guineafowl feather left by Wakusi with saliva on
the proximal end.

DISCUSSION
Wakusi carried the guineafowl feathers for at least
36 minutes. This may be an idiosyncratic behavior,although no similar behavior by her had previously been
observed, despite the fact that she has been identified andobserved for more than two decades. Given that she
picked up the feathers from the ground at least twice, it
could be said that she had an intention to collect and keepthem for a while. However, it is difficult to know why she
did so.
It is unlikely that Wakusi assumed that the feathers
represented bird meat. A guineafowl egg was once
observed to be eaten at Mahale, and other terrestrial birds
such as francolins and domestic chickens were alsocaught and eaten several times6. Therefore, it is possiblethat the chimpanzees consider guineafowl as prey.
However, when I observed chimpanzees eating other bird
species, they first discarded the feathers, and none
showed interest in the discarded feathers, although plentyof feathers were available. Therefore, it is unlikely thatchimpanzees show interest in feathers for this reason. In
addition, Wakusi never sniffed or licked the feathers.
The feathers did not seem to be like “toys,” either,because Wakusi did not show any attempt to manipulatethem in a playful manner, but just kept holding them inher hand, lips, or groin pocket. Although one adolescentmale showed slight interest, and one infant eventually

1. Nishida T, Kano T, Goodall J, McGrew WC, Nakamura M
1999. Ethogram and ethnography of Mahale chimpanzees.
Anthropol Sci 107: 141–188.
2. Nishida T 1973. The ant-gathering behaviour by the use oftools among wild chimpanzees of the Mahali Mountains. J
Hum Evol 2: 357–370.
3. Wrangham R, Peterson D 1996. Demonic Males: Apes andthe Origin of Human Violence. Houghton Mifflin, Boston.
4. Matsuzawa T 1997. The death of an infant chimpanzee at
Bossou, Guinea. Pan Afr News 4: 4–6.
5. Hayaki H 1985. Social play of juvenile and adolescent
chimpanzees in the Mahale Mountains National Park,
Tanzania. Primates 26: 343–360.
6. 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.
7. Appenzeller T 1998. Art: Evolution or revolution? Science
282: 1451–1454.

<NOTE>The Death of a Newborn
Chimpanzee at Mahale:
Reactions of its Mother andOther Individuals to the Body
Takanori Kooriyama
Laboratory of Wildlife Biology, Graduate School of Veterinary
Medicine, Hokkaido University

INTRODUCTION
Approximately 50% of infant chimpanzees in20
Mahale die before weaning1. The causes of preweaning
infant death include endemic disease, maternal
inexperience, poor nutrition, stillbirth, and infanticide.
Since Jane Goodall2 and Toshisada Nishida3 began
their long-term research on wild chimpanzees,
researchers have known that female chimpanzees oftentransport the bodies of their dead preweaned infants for aperiod extending from a couple of days to several months.
Matsuzawa observed a mother who was unable toabandon her 2-year-old daughter’s body and speculated
that this was because of her maternal affection4. Hosaka
et al. documented five cases of reactions to dead infantsby mothers and other chimpanzees5. One case impliedthat adult males would cannibalise the carcasses of newly
deceased infants. Another case described the longest
recorded transporting of a dead infant by its mother, atleast 3.5 months. A recent report from Mahale noted thatseven infant chimpanzees died in a flu-like epidemic in
2006, two of which were then carried by their mothers6.
Here I report another case of a chimpanzee mothercarrying the body of her dead newborn. I also describe thereactions of other chimpanzees to the body of the newborn.
Given the dearth of literature on this topic, this study
contributes to researchers’ understanding of the reactionsof chimpanzees faced with the death of a conspecific.

STUDY SITE AND CHIMPANZEES
Chimpanzees of M group in the Mahale Mountains
National Park, Tanzania, were studied. I studied Vera andher dead newborn male using the ad libitum observation
method. Other chimpanzees that interacted with the pair
were two young adult males (Cadmus and Primus), one
juvenile male (Teddy), one older adult female (Nkombo),
and one juvenile female (Carmen). Carmen was the
younger sister of Cadmus and had no younger siblings
herself.
OBSERVATIONSAt about 16:50 on 10 October 2008, as the core
party of M group reached Kansyana Research Station,
Vera appeared after 2 days’ absence carrying the body ofher firstborn around her neck. Vera had immigrated into
M group in 2006 and had seemed nulliparous. Thisdelivery was likely Vera’s first since joining the group,
which is consistent with past data from Mahale that
immigrant females usually need 2 years to conceive theirfirst infant1.
The infant may have been stillborn, because its head
seemed constricted at the nose point. At the constriction
point, the skin of the mouth was pressed toward the chin.
I hypothesised that this constriction was unlikely to haveresulted from a normal birth, but rather may haveoccurred while the infant was passing through the pelvic
canal. The infant had a red lesion on its head, but this did
not seem fatal. When Vera was found carrying the infanta couple of days after delivering it, the body had not been

Pan Africa News, 16(2), December 2009cannibalized; it smelled rotten but had no visible bite
marks. The infant was thought to be male because there
was no evidence of the characteristic sexual skin of afemale.
At first, Vera walked silently out of sight behind theothers with her dead infant. She was discovered by
Cadmus, who started making intimidating displays
toward her. Primus joined in. Cadmus then mounted Vera,
who fled, leaving the body.
At 16:55, Cadmus, Primus, Carmen, and Teddy sat
surrounding the body. Cadmus touched the body with his
index finger, lifted its leg to sniff the leg, anus, and penis
and widened the anus to inspect it. Vera tried several
times to retrieve the body but her efforts were in vain
because she could not approach Cadmus.
At 16:59, once Cadmus left, Carmen began carrying
the body in her mouth, followed by Teddy. After a minute,
Carmen climbed up a tree, carrying the body; here shetouched and pinched it around the anus. After 30 s, sheclimbed down and continued to carry the body, again
followed by Teddy. While Carmen was carrying the body,
Cadmus was nearby, watching the body.
At 17:03, when Carmen passed Cadmus with thebody, Cadmus started running to follow her. Carmen soon
stopped, at which point Cadmus caught up with her andsniffed the body. With Cadmus beside her, Carmen
started touching and pinching the skin and mouth of the
body and inspecting the anus, as Cadmus had done. She
also stepped several times. Cadmus left Carmen and the
body again.
At 17:06, Cadmus made an intimidating display andthus regained the body. He touched and sniffed it, just as
he had done before. Carmen sat beside Cadmus with her
hand on his shoulder, watching what he was doing.
Cadmus paused for a while and then left, leaving thebody.
At 17:07, Carmen retrieved the body and begantreating it violently, dragging it and hitting it against the
ground. She stepped and stomped on it repeatedly. As aresult, the surface hair of the body became damaged; it
was discoloured a dark green discoloration and gave off abad smell. Primus, who was nearby, did not touch the
body. Teddy followed Carmen and appeared to want achance to touch the body, but he was not successful.
Nkombo approached the body to sniff it but departed
soon afterward.
At 17:09, Carmen picked up the body and ran away
out of our sight. She reappeared with the others 30 min
later, without the body.

DISCUSSION
If my assumption that Vera carried her stillborn
infant is true, then this could be the first report onreactions to a stillborn infant by the mother and otherchimpanzees. However, it is possible that the partiallycannibalised newborn reported in Hosaka et al. couldPan Africa News, 16(2), December 200921
REFERENCES

Fig. 1. Cadmus peered into the dead body with his knuckles
on it. His sister, Carmen, laid her hand on her brother’s
shoulder.

have been stillborn, although the authors presumed that it
had been killed by some chimpanzee5. Roof et al.
reported that about 7.5% of the first parity of captive
chimpanzees result in stillbirths7. The rate of stillbirths
among Mahale chimpanzees is unknown, but it seems
natural that they should also deliver stillborn infants inthe wild.
My observations reveal that a female chimpanzee
can carry her dead infant even if insufficient time has
passed to establish a strong mother-infant relationship. It
is likely that Vera had carried the infant for 1 or 2 days
before being discovered. She might have continued tocarry the infant much longer had she not joined the otherchimpanzees, considering that she tried to regain
possession of the body several times.
Cadmus recognised the fact that the infant was deadand checked to see whether the body was fresh enough or
too rotten to eat. It is possible that he was motivated bycannibalism, because adult male chimps have awell-known tendency to eat dead infants5,8. Therefore, thebody might have been cannibalised by other chimpanzees
such as Cadmus if it had been found on the day of birth.
However, it is unclear why Cadmus did not eat it.
Possible explanations are putrefaction or some
psychological conflict.
Carmen showed similar reactions to the body as her
brother Cadmus. However, some of her reactions (e.g.,treating the body like a toy, a play partner, or a live infant)can be interpreted as imaginative play. In addition, Carmen
showed greater persistence for carrying the body than her
brother did, which can be explained by the tendency ofadolescent chimpanzees toward alloparental behaviour9.

ACKNOWLEDGEMENTS
This study was supported financially by the Ministry ofEnvironment (Japan) Global Environmental Research Fund
(F061) and by the Japan Society for the Promotion of Science
Global COE program. I thank TANAPA, TAWIRI, COSTECH,
and MMWRC for their cooperation throughout our fieldwork at
Mahale. I am also grateful to Dr. Kazuhiko Hosaka of Kamakura
Women’s University for his valuable comments.

1. Nishida T, Corp N, Hamai M, Hasegawa T,
Hiraiwa-Hasegawa M, Hosaka K, Hunt KD, Itoh N,
Kawanaka K, Matsumoto-Oda A, Mitani JC, Nakamura M,
Norikoshi K, Sakamaki T, Turner L, Uehara S, Zamma K
2003. Demography, female life history and reproductive
profiles among the chimpanzees of Mahale. Am J Primatol
59: 99–121.
2. van Lawick-Goodall J 1968: The behaviour of free-living
chimpanzees in the Gombe Stream Reserve. Anim Behav
Monogr 1: 161–311.
3. Nishida T 1973. The Children of the Mountain Spirits.
Chikuma-Shobo, Tokyo (in Japanese).
4. Matsuzawa T 1992. The death of an infant chimpanzee at
Bossou, Guinea. Pan Afr News 4: 4–6.
5. Hosaka K, Matsumoto-Oda A, Huffman MA, Kawanaka K
2000. Reactions to dead bodies of conspecifics by wild
chimpanzees in the Mahale Mountains, Tanzania. Primate
Res 16: 1–15 (in Japanese with an English summary).
6. Hanamura S, Kiyono M, Lukasik-Braum M, Mlengeya T,
Fujimoto M, Nakamura M, Nishida T 2008. Chimpanzee
deaths at Mahale caused by a flu-like disease. Primates 49:
77–80.
7. Roof KA, Hopkins WD, Izard MK, Hook Michelle,
Schapiro SJ 2005. Maternal age, parity, and reproductive
outcome in captive chimpanzees (Pan troglodytes) Am J
Primatol 67: 199–207.
8. Nishida T 1998. Deceptive tactic by an adult malechimpanzee to snatch a dead infant from its mother. Pan Afr
News 4: 4–6.
9. Nishida T 1983. Alloparental behavior in wild chimpanzeesof the Mahale Mountains, Tanzania. Folia Primatol 41:
1–33.

22

<FORUM>

Foreword
William C. McGrew
Dept. of Biological Anthropology, University of Cambridge, UK

Recent empirical studies of large carnivores andmammalian prey show their predator-prey relations to behighly malleable1. Moreover, much of the uncertainty
about the importance of predation by large felines onlarge-bodied apes stems from the rarity of observed
encounters between the two parties2. For example, Boesch3
asserts that most studied populations of chimpanzees
experience little or no such pressure, and so are not ‘real’
chimpanzees. Most data on the subject are indirect, so any
direct observations are to be prized4,5. The following note
was written in 1974 but never published; it is reproduced
here with only minor typographical corrections, in order to
add to this sparse data-set.

REFERENCES
1. Berger J 2008. The Better to Eat You With. Fear in the Animal
World. University of Chicago Press, Chicago.
2. Zuberbuehler K, Jenny D 2002. Leopard predation andprimate evolution. J Hum Evol 43: 8738–86.
3. Boesch C 2009. The Real Chimpanzee. Sex Strategies in theForest. Cambridge University Press, Cambridge.
4. Gandini G, Baldwin PJ 1978. An encounter between
chimpanzees and a leopard in Senegal. Carnivore 1: 107–109.
5. Hiraiwa-Hasegawa M, Byrne RW, Takasaki H, Byrne JM
1986. Aggression toward large carnivores by wild
chimpanzees of the Mahale Mountains National Park,
Tanzania. Folia Primatol 47: 8–13.

An Encounter between a Leopard
and a Group of Chimpanzees atGombe National ParkAnn H. Pierce
Primate Research Group, Department of Psychiatry and Behavioral
Sciences, Stanford University Medical Center, USA

INTRODUCTION
There has been considerable speculation concerning
the influence of predators on chimpanzee (Pan troglodytes)
populations1–3. This paper reports a complete observation of
an interaction between a leopard (Panthera pardus) and amixed group of chimpanzees and summarizes all previous
documented responses of chimpanzees during encounterswith leopards and lions (Panthera leo). These fieldobservations are currently limited to sites in western

Pan Africa News, 16(2), December 2009
Tanzania as follows:
1) In the Kasakati Basin, Izawa and Itani4,5 heard a single
young chimpanzee (7–8 years, sex uncertain) giving
loud calls. When they reached the site an hour later, thechimpanzee was shaking vines and still calling out as aleopard rested in a tree 25 m away. Upon noticing theobservers, the chimpanzee ran off.
2) In the Mahali Mountains, Nishida6 observed an adultmale chimpanzee silently throwing down twigs andshaking branches from a tree. The male descended to
the ground, and a leopard 40 m away moved into theundergrowth. Other chimpanzees were present in thearea.
3) In the Ugalla Hills, Kano7,8 found a group of
chimpanzees driven up trees by a young adult lion. They
were “screaming in a most unusual manner.” The lion
charged by and hit Kano, growled, then ran off.
4) van Lawick-Goodall9 reported two incidents at Gombe
National Park. Once a juvenile peered into a ravine andscreamed intermittently for 5 min. Its mother, who wascarrying a small infant, glanced down, then continued
feeding. Subsequent evidence showed that a leopard had
been present. On another occasion, a leopard vocalized
about 50 m from a group of chimpanzees feeding intrees. They made no apparent response.

OBSERVATIONSOn Nov. 18, 1974, I was alone observing six
chimpanzees from the Kahama Community, which range
in the south of Gombe National Park10,11. The groupconsisted of the following individuals:
Charlie (CH), adult male (approximate age, 26 years);
Willy Wally (WW), adult male with a partially paralyzedleg (ca. 26 years);
Sniff (SF), adult male (ca. 16 years);
Madam Bee (MB), old female with a paralyzed arm;
mother of LB and HB (ca. 35–40 years);
Little Bee (LB), nulliparous female with a “clubbed” foot;full estrous swelling at the time of the observation (ca.
14 years);
Honey Bee (HB), adolescent female with a small sexualswelling (10 years).
The chimpanzees left the nest site at sunrise (06.28 hr)
and began to walk south through woodland toward Kahama
Stream. HB and SF led as the group foraged along the
ground. A troop of redtail monkeys (Cercopithecus
ascanius), 350 m south of the nest site, was“alarm-chirping”12 from trees (5 trees, ca. 15 m high)surrounding a dense thicket (10–15 m diameter), in which afull-sized leopard of unknown sex was concealed.
At 06.52 HB climbed into a tree near the redtails,
stared at the leopard below, and “wraaahed” (long callindicating danger9). MB immediately climbed into the tree
near HB and began to wraaah too. LB ascended a nearby
tree. WW (who had been in close proximity to LB since
sunrise), proceeded southwest around the thicket, stoodwith hair erect, and peered into the undergrowth. SFglanced back with hair erect at CH, climbed onto a large
rock (2 m high, 10 m east of the thicket), and sat down
staring toward the thicket. MB and HB wraaahedpersistently from the tree above. CH scanned from the rear,

Pan Africa News, 16(2), December 2009
moved with hair erect slowly by the thicket, climbed
quickly up the rock, sat down in contact with SF, and faced
the concealed leopard.
At 06.54 the leopard rustled in the tangled vegetation
and appeared to lunge within the thicket toward SF and CH.
The chimpanzees wraaahed and “waa-barked” (loud,single-syllable bark given in threat9). The leopard’s
movements subsided. CH climbed down the rock, walked
with hair erect beside the thicket, and sat down by WW. CH
then stood bipedal within 10–15 m of the leopard andpeered into the vegetation. SF leapt with hair erect intovines near the females and stared at the leopard below.
Violently shaking vines for several seconds, SF wraaahed,charged away from the thicket, and sat down in the fork of alarge tree. Following this display, CH climbed silently to the
top of a slender tree nearby and the leopard left the thicket
headed south through dense ground cover.
HB stared at the leopard as it moved away, broke off abranch, and threw it into the thicket below. She descended
to the ground and followed the leopard at a distance of
15–20 m, with WW close behind. WW remained at the baseof the tree as HB charged up the trunk, slapping andstamping on the branches above the leopard. HB wraaahedand ran through the trees following the leopard as it
returned along the ground. During this pursuit, MB and LB
climbed through branches and stared toward the leopard
and HB. HB joined her family as the leopard passed
beneath them and lay down at the base of the tree.
At 07.00 the family stood close together, persistently
stamped on the branches, and wraaahed as the leopard
rubbed its forepaw against the ground. The males remained
silent and alert.
At 07.05 the leopard stood up and walked with a slight
limp toward riverine forest of Kahama Stream. The
chimpanzees descended, stood on the ground, stared at theleopard, and gave “hoos” (soft, low-pitched, single-syllable
sound often elicited in response to strange stimuli9). SF, HB,
and MB followed the leopard at a distance of 25 m, as it
moved out of sight.
At 07.10 CH directed a charging display at MB. She
stepped aside submissively as CH moved ahead. WW and
LB followed in the rear of the progression. At 07.12, CH,
travelling south with the others in the direction of theleopard, began to forage along the ground.

DISCUSSION
To my knowledge there is no direct evidence from any
study that large felines prey upon chimpanzees; however,the behavioral ecology of these predators has not beensystematically studied at any site. Furthermore, increasing
human activity may have altered the ranging patterns andreduced the number of large felines existing in these areas.
For example, leopards were considered common at Gombe
National Park only 15 years ago13. In the early 1960s, two
leopards ranged within Kakombe Valley, later the site of thebanana feeding station14. With the expansion of the Gombe
Stream Research Centre, leopards were no longer observedin the central valleys. In 1967, a small ranger station was
built 4 km south of the research centre on the lake.
Occasionally a leopard came down to the beach at night(pers. obs. and rangers), and in 1975, a leopard and its

23
offspring were observed in the nearby forest (Swai Saashiha,
pers. comm.). A small population of leopards apparently
exists in the southern area of the park. Two old male lions
once roamed within the reserve (Esilom Mpongo, pers.
comm.).
There is evidence from a trapping project that aleopard killed an infant chimpanzee in the Idambo region ofZaire in 196615. Rahm reported that her scouts brought afreshly-killed infant male aged about 5 mon to camp. His
whole lower abdomen had been torn open, the intestines
were visible, and the skin of his thighs and right forearm
were slashed. The scouts reported that after hearing noisesof fighting chimpanzees, they surprised a leopard attacking
a female carrying the infant. A large male chimpanzee in thegroup snatched the infant, carried it aside, then took a stick
and attacked the leopard. The men tried to carry the infant
away, but it cried and the male hurried back16. They
dropped the infant and the male returned to assist the female
with his stick. A fight had taken place at the described spot
when Rahm examined it, but there was no proof that aleopard had attacked the group. She states that the
chimpanzee’s wounds were inf1icted by claws and not bysnares and cables.
The leopard’s activity prior to detection by theadolescent female in the 1974 Gombe encounter is
unknown; however, it appears that the leopard attempted toprey upon the group. The following observed responses of
the chimpanzees indicate that leopards are regarded as
potentially dangerous: vigilance; orientation; movement
into close proximity with others; wraaah (alarm) calls,
primarily by the family of females; investigation; rock andtree climbing; displays from trees, including branch-shaking,stamping, and branch-throwing; and following the leopard
out of sight.
These behavioral patterns may communicate that theleopard is detected and induce it to move elsewhere insearch of food (see discussion of alarm signals17,18). The
movements of chimpanzees into close proximity withothers in potentially dangerous situations may function as adefense tactic, which affords protection to members of thesame community, such as related individuals, prospective
mating partners and males with whom there are strong
personal bonds (see discussion of male-malerelationships19).
Although mothers protect their offspring in a varietyof circumstances9, chimpanzees are also occasionallyobserved to respond in a protective manner to individuals
other than their own offspring, such as “adopting” anorphaned sibling9. In 1975, field assistants reported that anadolescent female encountered an unidentified snake, which
seemed to chase her. She climbed a tree and sat, watching
the snake, until her infant brother toddled along, seemingly
unaware of it. Suddenly, as the infant got close to the snake,the adolescent ran with hair erect down the tree, seized herbrother, and ran with him to their mother20. In Guinea aprime adult male ran down, picked up, and carried away animmature chimpanzee, who was unknowingly approaching
Nissen21.
Field assistants at Gombe National Park reported thatimmature and adult male chimpanzees carried around an
infanticide victim for hours after the fatal attack22. In 1974 adomestic dog charged toward the family described in the24leopard episode and bit MB on the head. As she slapped thedog with her arm, its tail was grabbed by LB and the dog
hurled aside and chased into nearby forest. HB stood
between them and watched (pers. obs.).
Prior to giving wraaah calls, the adolescent female
appeared alert to the alarm-chirps of the redtail monkey
troop above the leopard and climbed into a tree. Thisbehavior suggests the possible role of interspecificcommunication in detection of potential dangers23. The
adolescent also seemed to be the most highly aroused
member of the group in the leopard’s presence, which iscomparable to the responses of immature langurs (Presbytis
entellus) in encounters with leopards, a predator of thisspecies in Ceylon (Sri Lanka)17; the responses of juvenile
orang-utans (Pongo pygmaeus) to the presence ofhumans24; the stick-throwing response of an adolescentfemale, living with three other adolescent females and one
adult male, to a lion’s arrival on the island at Lion Country
Safari, California (Patrick McGinnis, pers. comm.); theresponses of juvenile Japanese monkeys (Macaca fuscata)
to novel objects25.
In Zaire and Guinea investigators have tested the“dehumanization hypothesis” of African ape evolution by
exposing large parties of savanna and forest chimpanzees toa stuffed leopard. Some of the open country chimpanzees
attacked the experimental object with sticks used as
clubbing tools2,8,26. Such behavior has not been observed inwestern Tanzanian chimpanzees during encounters withleopards in the wild and suggests that chimpanzees may
distinguish between and respond differently to live andstuffed leopards.

ACKNOWLEDGEMENTS
I am grateful to Dr. Jane Goodall for the opportunity to study
at Gombe National Park as a research assistant; the Tanzania
National Park personnel, particularly the Director, Derek Bryceson,for their cooperation; the W.T. Grant Foundation, which financed
the research; and the Stanford University Primate Research Group,particularly Dr. David Hamburg, for providing a stimulating
environment in which to analyze data. I also thank C. Busse, R.
Wrangham, W.J. Hamilton III, J. Silk, P. Lee, G. Heidrich, E.v.Z.
Bergmann, S. Smith, C. Czeis1er, J. Goodall, D. Hamburg, andothers for their invaluable advice and critiques in the preparation ofthis manscript. Finally, my thanks to E. Moore, E. Mpongo, and C.
Chiwaga for their active assistance in searching for and following
the southern chimpanzees.

REFERENCES
1. Chance M, Jolly C 1970. Social Groups of Monkeys, Apes, andMen. Dutton, New York.
2. Kortlandt A, Kooij M 1963. Protohominid behaviour inprimates. Symp Zool Soc London 10: 61–88.
3. van Lawick-Goodall J 1970. Tool-using in primates and othervertebrates. In: Advances in the Study of Behavior 3, Lehrman
DS, Hinde RA, Shaw E (eds), Academic Press, New York, pp.
195–249.
4. Itani J 1970. Chasing Wild Chimpanzees. Chikuma-Shobo,
Tokyo (in Japanese).
5. Izawa K, Itani J 1966. Chimpanzees of Kasakati Basin,
Tanzania. Kyoto Univ Afr Stud 1: 73–156.
6. Nishida T 1968. The social group of wild chimpanzees in the
Mahali mountains. Primates 9: 167–201

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7. Kano T 1972. The distribution and adaptation of thechimpanzee on the eastern shore of Lake Tanganyika. Kyoto
Univ Afr Stud 7: 37–129.
8. Kortlandt A 1972. New Perspectives on Ape and Human
Evolution. Stichting voor Psychobiologie, Amsterdam.
9. van Lawick-Goodall J 1968. The behaviour of free-living
chimpanzees in the Gombe Stream Reserve. Anim Behav
Monogr 1: 161–311.
10. van Lawick-Goodall J 1975. The behavior of the chimpanzee.
In: Hominisation und Verhalten, Kurth F, Eibl-Eibesfeldt I(eds), Gustav Fisher, Stuttgart, pp. 72–135.
11. Wrangham R 1975. Behavioura1 Ecology of Chimpanzees inGombe National Park, Tanzania. PhD thesis, University ofCambridge, Cambridge.
12. Marler P 1973. A comparison of vocalizations of red-tailed
monkeys and blue monkeys, Cercopithecus ascanius and C.
mitis, in Uganda. Z Tierpsychol 33: 223–247.
13. Thomas DK 1961. The Gombe Stream Game Reserve.
Tanganyika Notes Records, 56: 35–39.
14. van Lawick-Gooda11 J 1967. My Friends the Wild
Chimpanzees. National Geographic Society, Washington, D.C.
15. Rahm U 1967. Observations during chimpanzee captures in
the Congo. In: Progress in Primatology, Gustav Fischer,
Stuttgart, pp. 195–207.
16. Rahm U 1976. Personal communication by letter dated May 2,
1976.
17. Eisenberg JF, Lockhardt J 1972. An ecological reconnaissanceof Wilpattu National Park, Ceylon. Smithsonian Contrib Zool
101: 1–118.
18. Smythe N 1970. On the existence of “pursuit invitation”
signals in mammals. Am Nat 104: 491–494.
19. Bygott D 1974. Agonistic Behaviour and Dominance of Wild
Chimpanzees. PhD thesis. University of Cambridge,
Cambridge.
20. Goodall J (ed) 1975. Gombe Journal: Part 3. Gombe Stream
Research Centre Reports. Unpublished manuscript.
21. Nissen HW 1931. A field study of the chimpanzee:
Observations of chimpanzee behavior and environment inwestern French Guinea. Comp Psychol Monogr 8: 1–122.
22. Goodall J, Bandora A, Bergmann E, Busse C, Matama H,
Mpongo E, Pierce A, Riss D 1979. Inter-community
interactions in the chimpanzee population of the Gombe
National Park. In: The Great Apes, Hamburg DA, McCown
ER (eds), Benjamin/Cummings, Menlo Park, pp. 13–53.
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I (ed), Holt, Rinehart, and Winston, New York, pp. 544–584.
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Piper, Muenchen.
Pan Africa News, Vol. 16, No.2
Published in December, 2009
Address:

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