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JUNE 2010
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

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VOL. 17, NO. 1

Contents<NOTE>
Case study of an unusual human–chimpanzee conflict at
Bulindi, Uganda
Matthew McLennan

1<NOTE>
New way to dip for honey is the first observation of tool-use
in wild chimpanzees of Nyungwe National Park, Rwanda
Julian Easton

5<NOTE>
Bird in the hand: Bossou chimpanzees (Pan troglodytes)
capture West African wood-owls (Ciccaba woodfordi) but not
to eatSusana Carvalho, Yumi Yamanashi, Gen Yamakoshi,& Tetsuro Matsuzawa

6<NEWS>
Puffy inherits a habit from her father?
Toshisada Nishida

10<NOTE>

Case Study of an Unusual
Human–Chimpanzee
Conflict at Bulindi, Uganda
Matthew McLennan
Department of Anthropology & Geography, Oxford Brookes
University, UK(E-mail: mmclennan@brookes.ac.uk)

INTRODUCTION
Across equatorial Africa expanding humanpopulations and forest clearance for agriculture has meantthat increasingly humans and great apes occupy the same
habitat. Interactions between apes and people in sharedenvironments are often characterized by competition andconflict1–6. In this report I describe an unusual
human–ape conflict involving a newly-studied
community of ≥ 25 chimpanzees (Pan troglodytes
schweinfurthii) in heavily disturbed forest–farm habitat at

2

Pan Africa News, 17(1), June 2010

Bulindi, Western Uganda.

Study Site
Bulindi is situated between 1°27’−1°30’N and
31°26−31°30’E, 25 km south of Budongo Forest. The
chimpanzees’ territory comprises a network of small (≤50 ha) riverine forests within the vicinity of 11 permanentvillages. These unprotected forests are being logged fortimber and cleared for cultivation3. The chimpanzees
regularly consume agricultural foods. They are nothunted but experience high levels of harassment fromvillagers. Although familiar with people, the apes are
unhabituated6.

CONFLICT CASE STUDY
BackgroundOf five main riverine forests at Bulindi, Kaawango
forest in the northeast was least utilized by chimpanzees
during this study (Oct 2006–Jan 2008). No evidence

Fig. 1. Map showing riverine forest fragments at Bulindi; the surrounding matrix is dominated by farmland. Italicized names indicate
forests mentioned in the text. The location of the homestead visited by chimpanzees is indicated, bordering Kaawango forest. The
chimpanzees’ core range extends to the southwest.

Pan Africa News, 17(1), June 2010
indicated chimpanzees visited Kaawango between
October 2006 and April 2007. However, on 15th May,
chimpanzees raided mangos at a home bordering theforest (Fig. 1). They returned to this homestead for
mangos ≥ 5 times in May–June, after which the fruits
were gone. On each occasion they travelled to Kaawango
from Kiseeta or Kyamalera forests, in the core of theirrange (Fig. 1), apparently specifically for these mangos,
which had fruited earlier than other mango trees in theirterritory. They spent little time within Kaawango forest.
Unlike most village homes, this house had concrete
walls and glass windows. Ornamental shrubs and trees
were planted around a grass lawn. The owner, a senior
government official, resided in the capital Kampala but
occasionally visited Bulindi. Metal doors on one side of
the house had glass window panes in which chimpanzees
could see their reflections, and this ‘mirror’ provoked
considerable interest and excitement in the apes. Two
field assistants (FA) to M.M. lived near this homestead,and much of the following account of chimpanzee
behavior at the homestead is based on their observations.
15th May. Chimpanzees crossed to Kaawango from
Kiseeta and raided mangos. In mid-afternoon, two FAs
arrived at the homestead to find three adult malechimpanzees by the door. One was swaggering bipedally
in front of the glass. Local residents wanted to chase themaway with dogs but the FAs prevented them from doing
so. When M.M. arrived, the chimpanzees were in nearby
trees. Feces were found around the homestead.
17th May. Four adult males raided mangos late in
the afternoon. Two FAs found them displaying vigorouslyon the lawn. They had broken several branches fromsmall trees; one male brandished a branch while
swaggering in front of the glass. Another male slowly atea mango whilst staring intently at the glass. They paid
little attention to FAs, who watched from a distance.
23nd May. At 0630, a FA found an adult male, an
estrous female and a juvenile at the homestead. The male
was displaying at the glass door. The apes left upondetecting the FA.
28th May. Four adult males raided mangos at 1915
and nested in trees immediately behind the house. At
0645 the next morning, a FA found them on the lawn infront of the door. They had again broken branches from
ornamental trees and were examining the window. Before
0700, the males had left for Kiseeta, crossing the mainroad at the trading centre.
8th June. Three adult males raided the last of themangos at dusk, nested nearby, and returned to thehomestead at first light. Again, they displayed on thelawn, but left when a FA approached.
18th July. At around noon, school children saw
chimpanzees ‘dancing’ at the homestead. When wearrived at 1600, five adult males were resting in trees
behind the house. They crossed to Kiseeta shortly
afterwards. Feces and broken branches were strewn

3
across the lawn.
12th August. Before 1130, five adult males had
arrived at the homestead. It was Sunday and local
residents and school children joined FAs to watch themfrom a distance. The males displayed all around thecompound, breaking branches from trees and dragging
them. Although people’s presence may have exacerbatedtheir excitement, the apes did not threaten them directly.
They appeared to want to get inside the door. One maleflailed a branch against it, breaking an upper pane ofglass. Another was seen trying to push a stick between
the doors in an apparent attempt to prise them apart(woody particles were subsequently visible in the narrowgap separating the door’s two sides). In between bouts ofdisplaying, the males groomed or rested, eventuallyleaving for Kiseeta before 1530. In addition to the broken
pane, the metal panelling beneath the glass was heavily
scratched by the apes’ fingernails.
The latter two visits were made after mango season
was over. Both times, the chimpanzees spent several
hours around the homestead, apparently without feeding,
before returning to other forests in their core range.

Confrontation between Chimpanzees and the House
Owner

Shortly after 1300 on 17th August, a FA telephoned
M.M in Kampala, informing him of a “big fight” between
chimpanzees and the homestead’s owner, who had
arrived the previous day. The owner was apparently
shooting at the apes, which were in a frenzied state intrees behind the house. The owner was also highly
agitated and could not be approached. It was a prolonged
confrontation; shots were still being fired approximately
30 minutes later. According to FAs, chimpanzees
remained at the site calling and drumming until lateafternoon (i.e. several hours). The following morning,FAs searched the forest but found no dead apes and no
blood.
On 22nd August M.M. visited the owner who gave
the following account of the incident: at around 1300 or
earlier, he was inside his house and heard loud banging
against the door. Two chimpanzees were outside,
immediately in front of the glass. He opened gates at theside of the house, which open to a small courtyard, andwent to chase them away. The animals charged towards
him, vocalizing loudly; one was carrying a branch.
Shocked, he ran back into the courtyard, shutting thegates. He was convinced the apes wanted to physicallyattack him and began shooting with his rifle in the air tofrighten them away. The chimpanzees ran 50m to
Eucalyptus trees and, still vocalizing, broke branches tothreaten him. (According to the FA, other apes were indense undergrowth nearby, vocalizing and drumming).
The owner continued firing in the air. After the initialconfrontation, chimpanzees twice returned to the4homestead; each time, the owner fired shots to repel them.
He said he did not shoot them because he knew
chimpanzees are legally protected. The six identified
adult males at Bulindi were all subsequently seen by the
research team in good health.
On the afternoon of this meeting, a mixed party of ≥9 chimpanzees travelled to Kaawango. FAs found five
adult males on the lawn by the glass door. At theirapproach, one male displayed towards them dragging abranch. However, after several minutes, the ownerarrived, having heard chimpanzees were at his house, anddrove his car at the apes causing them to flee. He
subsequently agreed to cover the lower glass panes of thedoor with cloth as a temporary solution. During theremaining five months of research, chimpanzees travelled
to Kaawango ≥ 8 times and raided papaya, sugarcane,cocoa and oranges in villages surrounding the forest.
They visited the homestead ≥ 5 times. On three occasions,adult males were discovered seated calmly by the door,
now covered with cloth; each time, they returned to theforest at our approach. In October, chimpanzees weretwice chased away by the owner, apparently without
incident.

DISCUSSION
Captive chimpanzees are famously capable ofself-recognition in mirrors7. However, the capacity for
self-recognition is most pronounced in subadult animals(8–15 yrs), and the proportion of subjects showing
self-recognition declines sharply among adults8. In thisanecdotal report, wild adult male chimpanzees at Bulindi
were evidently fascinated by their reflections in the glass
door of a house, returning to the home once mangos – theinitial attractant – had finished fruiting. The chimpanzees’
range includes many streams as well as papyrus swamps,
and after heavy rains stretches of riverine forest become
flooded. Consequently, the apes have ample opportunity
to see their reflections in water. Young chimpanzees at
Mahale were observed examining their reflections inpools and streams9, but it is not yet known if similar
behavior occurs at Bulindi. Although the males at times
intently examined the door, their high state of arousal
implies that they most likely saw ‘other’ chimpanzees
behind the glass. The house is situated outside of theircore area in the northeast corner of the chimpanzees’
known range. Visiting parties usually comprised adultmales only. This raises the possibility that the males had
identified the homestead as a location for territorial
encounters with extra-group strangers. A motivation to
engage in territorial confrontation at the homestead isperhaps reflected in their willingness to remain at theconflict scene to monitor and threaten the owner, despite
the noisy gunshots. After the windows were covered,chimpanzees made occasional visits to the house, but
their behavior was no longer excitable and aggressive.
With increasing human penetration into great ape
habitats and growing interest in the conservation

Pan Africa News, 17(1), June 2010
implications of human–great ape interactions inhuman-dominated landscapes4–6, further reports of novel
human–ape conflicts are anticipated.

ACKNOWLEDGEMENTS
Research permission was granted by the President’s
Office, the Uganda National Council for Science andTechnology, and the Uganda Wildlife Authority. I thankDan Balemesa, Gerald Sunday Mayanda, Tom Sabiiti,
and Moses Ssemehunge for assistance in the field. The
research was funded by an ESRC/NERC interdisciplinary
studentship to M.M., and a Leverhulme Trust award to
Catherine Hill, who provided helpful comments on the
manuscript.

REFERENCES
1. Reynolds V 2005. The Chimpanzees of the Budongo Forest.
Oxford University Press, Oxford.
2. Madden F 2006. Gorillas in the garden: human–wildlife
conflict at Bwindi Impenetrable National Park. Policy Matters
14: 180–190.
3. McLennan MR 2008. Beleaguered chimpanzees in the
agricultural District of Hoima, Western Uganda. Primate
Conserv 23: 45–54.
4. Hockings KJ 2009. Living at the interface:
human–chimpanzee competition, coexistence and conflictin Africa. Interact Stud 10: 183–205.
5. Hockings K, Humle T 2009. Best Practice Guidelines for
the Prevention and Mitigation of Conflict Between Humans
and Great Apes. IUCN/SSC Primate Specialist Group.
Gland, Switzerland.
6. McLennan MR, Hill CM 2010. Chimpanzee responses toresearchers in a disturbed forest–farm mosaic at Bulindi,
Western Uganda. Am J Primatol (in press)
7. Gallup Jr GG 1970. Chimpanzees: self recognition. Science
167: 86–87.
8. Povinelli DJ, Rulf AB, Landau KR, Bierschwale DT 1993.
Self-recognition in chimpanzees (Pan troglodytes):
distribution, ontogeny, and patterns of emergence. J Comp
Psychol 107: 347–372.
9. Nishida T, Matsusaka T, McGrew WC 2009. Emergence,
propagation or disappearance of novel behavioral patterns in
the habituated chimpanzees of Mahale: a review. Primates
50: 23–36.

Pan Africa News, 17(1), June 2010

5

<NOTE>New Way to Dip for Honey is theFirst Observation of Tool-use in
Wild Chimpanzees of Nyungwe
National Park, Rwanda
Julian Easton
Nyungwe Forest Conservation Project (P.C.F.N.) / W.C.S. Rwanda(E-mail: juleseaston@yahoo.co.uk)

Honey dipping is well documented in wild
chimpanzee populations across their range1,2. Nyungwe
National Park in the south-western region of Rwanda has
anestimatedchimpanzee(Pantroglodytes
schweinfurthii) population of 340 individuals3, and thisrecord is the first documented report of tool use from thissite. There are unconfirmed reports from park field staff
of chimpanzees dipping for honey with sticks and there
have been several unconfirmed observations of discarded
tools left protruding from tree hollows. Theseobservations suggest honey-dipping is one type oftool-use practiced by this chimpanzee population.
However I made a new observation of a novel way of
manipulating a honey-probe tool by a wild chimpanzee,
which expands the breadth of tool use of wild populations
as a whole. I made a serendipitous observation of atool-use episode in which an adult male chimpanzee
probed for honey by manipulating the tool firstly with his
hand and then by clamping the honey-probe stick
between his teeth using his jaw muscles and manipulating
it with his mouth. In October 2009, the individual
ascended a tree with a long stick of approximately 80cm
in length and then stripped the branch of bark and leaves
with his teeth. Using his right hand, he forced the stick
into a beehive (Possibly Apis mellifera), in a hollow inthe trunk of the tree. However, he was unable to gain
sufficient force to break into the hive, and so repositioned
himself upside down and took the stick in his mouth. He
gripped the stick with his molar and incisor teeth. It was
positioned in such a way that it passed on the inside of his
upper and lower right canines, and then extended
backwards out of the side of his mouth past his rightcheek and forwards from between his incisors (see Fig. 1).
With the stick firmly gripped between his teeth, he used
his arms, body weight and neck muscles to push his headtowards the tree and at the same moment force the tool,
gripped in his mouth, into the bee hive in order to breakinto it. After re-positioning himself and attempting for afurther five attempts, he successfully broke into the hive.
Once the probe had penetrated the hive, he released the
stick from his mouth and manipulated it with his hand.
He then removed the probe from the hole using his handFig. 1. Chimpanzee probing for honey using a newtechnique, the jaw–clamp.

and wiped the visible honey from the tip with his lips. He
repeated probing for circa. 35 minutes in this way by
forcing probes into the hollow both by using the tool in
his hand and also clamping the tool with his teeth when
he required more leverage. When the first honey probe
snapped, he replaced with another, using a total of fourhoney-probes during this single tool-using episode. All
subsequent honey-probes were made by breaking off
small branches of the tree in the vicinity of the bee hive
before stripping them of bark and leaves with his teeth.
All honey-probes were manipulated both by hand and by
his mouth. This observation provides the first descriptionof such a “jaw–clamp” manipulation in the handling oftools in free-ranging chimpanzees.
This observation expands the diversity in which
tools are manipulated by chimpanzees and may provide
an alternative method to pounding beehives using honeyhammers / clubs which has been documented in othersites in Central Africa to access beehives, but has not yet
been documented in Nyungwe4-6. There are records of wildorangutans using their mouths to manipulate tools, in which
they clenched probes in their teeth to extract food fromholes7 and placed leaves next to their mouth in order toamplify sounds that they make8.
This opportunistic observation provides both thefirst documentation of any tool-use in this chimpanzee
population and also expands the diverse set of tool-use
abilities in free-living chimpanzees. Preservation of
distinct tool-using behaviours is an importantconsideration for conservation strategies and this6

Pan Africa News, 17(1), June 2010

observation highlights the need for further research of thechimpanzee population living in Nyungwe National Park,
Rwanda.

4. Bermejo M, Illera G 1999. Tool-set for termite-fishing andhoney extraction by wild chimpanzees in the Lossi Forest,
Congo. Primates 40(4): 619–627.

REFERENCES

5. Hicks TC, Fouts RS, Fouts DH 2005. Chimpanzee (Pantroglodytes troglodytes) tool use in the Ngotto Forest, Central
African Republic. Am J Primatol 65: 221–237.

1. McGrew W 1994. Tools compared, the material culture. In:
Chimpanzee Cultures, Wrangham RW, Mcgrew WC, de Waal
FBM, Heltne PG (eds), Chicago Academy of Sciences, pp.
162-166.
2. Sanz C, Morgan DB 2009. Flexible and persistent tool-using
strategies in honey-gathering by wild chimpanzees. Int J
Primatol 30: 411–427.
3. Chao N, Mulindahabi F, Ntare N, Easton J, Plumptre A,
Ndikubwimana I, Rugyerinyange L 2010. Biodiversity survey
of Nyungwe national park, Rwanda (2009). Technical Reportfor Rwanda Development Board. WCS/ PAB/ GEF/ UNDP.

6. Sanz C, Morgan DB 2007. Chimpanzee tool technology in theGoualougo Triangle, Republic of Congo. J Human Evol 52:
420–433.
7. O’Malley RC, McGrew WC 2000. Oral tool use by captive
orangutans. Folia Primatol 71: 334-341.
8. Van Schaik CP, Ancrenaz M, Borgen W, Galdikas B, Knott CD,
Singleton I, Suzuki A, Utami SS, Merrill M 2003. Orangutan
cultures and the evolution of material culture. Science 299:
102-105.

<NOTE>

Bird in the Hand: Bossou Chimpanzees (Pan troglodytes)
Capture West African Wood-owls (Ciccaba woodfordi) but Not
to EatSusana Carvalho 1, Yumi Yamanashi 2, Gen Yamakoshi 3 & Tetsuro Matsuzawa 2
1. Leverhulme Centre for Human Evolutionary Studies, University of Cambridge, UK / Centre of Research in Anthropology and Health, University of
Coimbra, Portugal (E-mail: scr50@cam.ac.uk)
2. Primate Research Institute, Kyoto University, Japan
3. Centre for Asian and African Area Studies, Kyoto University, Japan

Fig. 1. Adolescent male PL after capture of the owl, carrying it in right hand.

Pan Africa News, 17(1), June 2010
INTRODUCTION
Chimpanzees rarely capture an animal without
eating it, but this puzzling pattern occurs in both natureand captivity, mostly by youngsters1-6. At Bossou in
Guinea, chimpanzees rarely hunt for meat; their homerange presents few suitable prey7. From 1976 to 2008,
only one bird (West African wood-owl, Ciccaba
woodfordi) capture was recorded, and it was consumed7.
In 2009, we saw two captures of this species, which were
not followed by consumption. Instead, both raptors wereused as toys, as previously observed at Bossou with tree
hyrax, Dendrohyrax dorsalis3. Here we describe thecaptures and discuss the implications of this behavior
from an evolutionary perspective.

7
shrub forest. The chimpanzees are fully habituated tohuman observers. The bird captures occurred between
November 2008 and November 2009, during (mostly)
nest-to-nest follows totaling 690 hours. These combined
focal and scan sampling, and we videotaped andphotographed events to supplement direct observations.
Rare or unusual events were recorded ad libitum. Bothepisodes were at nearly the same location, on the eastern
slope of Guein Hill (N 7.38.698; W 8.30.102).

RESULTS

Case 1: On 21 February 2009, S.C., G.Y. and two fieldassistants (J.D., G.Z.) followed a party of sevenindividuals (JR, JY, PL, JJ, TA, FF, YL), eating fruits. At
0941hr, we heard the alarm call of a bird nearby, about
20m away. Upon approaching the site, we saw an adultowl swooping over the head of an 11-year-old adolescent
METHODS
male chimpanzee (PL), apparently trying to “mob” him.
Site and Subjects
PL descended a short distance, carrying in his mouth a
Bossou is a long-term chimpanzee (Pan troglodytes
motionless owl, apparently dead, and then placed theverus) study-site in Africa8. Its chimpanzees range overchick in a groin pocket. He had feathers in his mouth.
15 km2, dominated by primary and secondary forest andFrom 0945hr to 1021hr: PL made a tree nest, then moved
surrounded by cultivated fields, abandoned fields andto another tree nearby and made two large tree nests, lay
down in each, and rested
supinely with the bird on his
belly. At 1027hr PL tapped
his feet on the nest andfeathers fell to the ground.
Two minutes later, he left thethird nest, but then returned toenlarge it and began to playwith the carcass, balancing it
on his feet while lying supine.
At 1034hr, he again left thenest, carrying the owl in byhand (Fig. 1). From 1037hr to1102hr: PL moved to anothertree and carrying the owl in
his groin pocket, while eating
fruits. He built a fourth nest at
1044hr, where he again
played with the owl,balancing the carcass on his
arm (Fig. 2) and grooming it.
He left the nest anddescended carrying the bird
(now featherless) (Fig. 3).
From 1102hr to 1229hr: On
the ground PL carried thecarcass dorsally, on hisshoulder, while eating leaves.
He inspected the dead bird,
touched it with index finger
and put the finger into his
Fig. 2. Playing in large day nest, balancing dead bird with one hand, while lying supine.
mouth, then moved away toeat fruits. He then rested on

8

Pan Africa News, 17(1), June 2010

Fig. 3. Balancing the owl, after plucking its feathers.

Fig. 4. End of event 1: “toy” broken in pieces and PL is resting after 4hr of play.

the ground, with the owl lying on his belly; in thisposition, he touched alternately his genitals and the owl
with his hand. Next, PL made a ground nest and balanced
the carcass with both hands while lying supine. From
1229hr to 1346hr: PL pulled off the bird’s wings. He
enlarged his ground nest and rested, whisking away flies
circling the carcass (Fig. 4). At 1346hr PL emerged fromhis ground nest, leaving the carcass behind. No otherchimpanzees showed interest in PL’s activities throughout
the events.

Case 2: On 22 October 2009, Y.Y. and a field assistant(J.D.) followed PL in the Guein area. At 0855hr, theysuspended observations for about an hour, then at 1003hr,
resumed focal sampling on him. He sat on the groundnext to the alpha male, holding a dead owl in his hand.
From 1008hr to 1011hr: PL travelled and fed carrying thebird in his hand. An infant male (FL) approached him andpeered closely, as PL groomed the carcass. After the
infant departed, PL continued examining the bird forPan Africa News, 17(1), June 2010
about 10 seconds, then abandoned it. The researcher
immediately collected the carcass and verified thatfeathers had been plucked and part of the viscera had
been extracted.

DISCUSSION
These are the first observed bird captures at Bossou
not followed by prey consumption. Nishida et al.9 noted
at Mahale, Tanzania, that parts of the carcass sometimes
were abandoned without consumption. As at Bossou,
these events involved young solitary individuals who
opportunistically encountered prey by chance. Bonobos(Pan paniscus) showed similar exploration and grooming
after capturing infant monkeys, which were handled likedolls10.
Our observations echo previous reports from
Bossou: Prey was found by chance, without active pursuit,and adults showed little or no interest in the prey7.
Bossou’s chimpanzees eat few vertebrate prey11,
including only one species of bird, this one. Owls may berelatively easy to catch, as they roost during the day in the
canopy. A recent report from Mahale12, on some of thevery few bird predations by chimpanzees, differs
somewhat from the Bossou events, for example, birds at
Mahale are captured and consumed. However, several
behavioral similarities emerge: catching of birds issolitary and opportunistic, does not require group hunting,the targeted birds are nestlings, and idiosyncrasies may
explain why some chimpanzees tend to repeat thisbehavior.
Availability (or lack) of resources and distribution
and abundance of prey may explain the lack ofemergence of pursuit-hunting at Bossou. Or, some of the
behavioral patterns seen rarely at Bossou, or seen only inthe early years of research, may not have spread or
persisted, due to the migration or disappearance of key
members of this small group, or to too few potential extra
innovators arriving through immigration. If prey-captureis sporadic and unpredictable, opportunities to watch ahunter are rare, and so observational learning and culturaltransmission is hindered. Further research is needed toassess systematically the abundance and availability ofprey and of learning opportunities.

ACKNOWLEDGEMENTS
DNRST and IREB (Guinea); MEXT-16002001,
JSPS-ITP-HOPE, JSPS-gCOE-Biodiversity, and F-06-61
of the Ministry of Environment, Japan; Mitsui & Co., Ltd.
Environment Fund; FCT-Portugal;The Wenner-Gren
Foundation; Luís Da Silva for exceptional illustrations;Field assistants; S. Assis, D. Biro, T. Humle, W. McGrew,
for comments on the manuscript.

REFERENCES
1. Teleki G 1973. Notes on chimpanzee interactions with small
carnivores in Gombe National Park, Tanzania. Primates 14:9407-412.
2. Goodall J 1986. The Chimpanzees of Gombe: Patterns of
Behaviour. Cambridge, MA, Harvard University Press.
3. Hirata S, Yamakoshi G, Fujita S, Ohashi G, Matsuzawa T
2001. Capturing and toying with hyraxes (Dendrohyrax
dorsalis) by wild chimpanzees (Pan troglodytes) at Bossou,
Guinea. Am J Primatol 53: 93-97.
4. Goldberg TL, Gillespie TR, Rwego IB, Kaganzi C 2006.
Killing of a pearl-spotted owlet (Glaucidium perlatum) by
male red colobus monkeys (Procolobus tephrosceles) in aforest fragment near Kibale National Park, Uganda. Am J
Primatol 68: 1007-1011.
5. Videan E, Fritz J, Murphy J 2007. Hunting and occasional
consumption of prey items by chimpanzees at the Primate
Foundation of Arizona. Int J Primatol 28: 477-481.
6. Ross SR, Holmes AN, Lonsdorf EV 2009. Interactions
between zoo-housed great apes and local wildlife. Am J
Primatol 71: 458-465.
7. Sugiyama Y 1989. Description of some characteristic
behaviors and discussion on their propagation process
among chimpanzees of Bossou, Guinea. In Behavioral
studies of Wild Chimpanzees at Bossou, Guinea (Sugiyama,Y., ed.), pp. 43-47. Inuyama: Kyoto University Primate
Research Institute.
8. Sugiyama Y, Koman J 1979. Social structure and dynamics
of wild chimpanzees at Bossou, Guinea. Primates 20:
323-339.
9. Nishida T, Uehara S, Nyundo R 1979. Predatory behavior
among wild chimpanzees of the Mahale Mountains.
Primates 20: 1-20.
10. Sabater Pi G, Bermejo M, Illera G, Vea JJ 1993. Behaviourof Bonobos (Pan paniscus) following their capture ofmonkeys in Zaire. Int J Primatol 14: 797-804.
11. Sugiyama Y, Koman J 1987. A preliminary list of
chimpanzees' alimentation at Bossou, Guinea. Primates 28:
133-147.
12. Fujimoto M, Shimada M 2008. Newly observed predationof wild birds by M-group chimpanzees (Pan troglodytes
schweinfurthii) at Mahale, Tanzania. Pan Afr News 15:
23-26.

10

Pan Africa News, 17(1), June 2010

<NEWS>

Puffy Inherits a Habit fromHer Father?
Toshisada Nishida
Japan Monkey Centre, Japan(E-mail: nishida.mahale@gmail.com)

In August 2009, I visited Mahale after a two-year
absence and stayed for two weeks. My field season
was so short that I made every effort to observe as
many chimpanzees as possible to catch a glimpse of
behavioral changes among M group chimpanzees.
A nine-year-old female, Puffy, was one of thechimpanzees I wanted to observe most because she
had lost her mother, Pinky, during the outbreak ofrespiratory disease in 20061.
Gwekulo, a sterile female, was interested in
caring for infants of other females or orphaned infants.
Thus, she adopted a 4-year-old female orphan Pipi in1994 after Pipi’s mother died of another respiratory
outbreak in 19932, although she was also interested ina two-year-old male infant of Wakampompo. By
September 1995, Gwekulo had begun to carry Primus,
Puffy’s elder brother, 4 years old at that time. Since
then, Gwekulo had become Pinky’s great partner.
Pinky gave birth to Puffy in 2000 long after Primus
was weaned, and Gwekulo became Puffy’s earnest
babysitter. Gwekulo and Pinky not only became closefriends but also formed a formidable coalition. Anygroup member in conflict with either of them had tofight both of them, and Pinky seemed to be one of themost dominant females of the M group.
After Pinky’s death, Gwekulo continued to carefor orphaned Puffy. However, after Puffy became anadolescent, Gwekulo’s keen “maternal” drive wasdiverted from Puffy and directed more to younger
infants although she continued to groom Puffy. In
August 2009, Gwekulo was most interested in Xtina’s
one-year-old infant and cared for it wheneveropportunity allowed. This baby had at least threebabysitters: Puffy and the baby’s elder sister,nine-year-old Xantippe in addition to Gwekulo. Theytransported and groomed the baby one after another.
Xtina seemed to enjoy traveling without carrying acharge during many of the daytime hours.
On August 25, I was surprised to watch Puffy’s
behavior. When she pant-grunted to a young adultmale, Orion, she touched her right nipple with her left
fingers. This was the first time I observed her do so.
Puffy massaged her left nipple with her right thumband then pant-hooted for 5 seconds. She again
continued this self-massage for 30 seconds (Fig. 1).
I was shocked because the way she fumbled thenipple was so much like that shown by an adult male,
Alofu. He has been seen to massage his nipple since
he was an adolescent male3,4. He was notorious for thishabit5 because he seemed to show this “nipple press”
or “fumble nipple”6 whenever he appeared to beuneasy, e.g., when he pant-grunted to the alpha male orwhen he anticipated the approach of aggressive males.
Alofu’s nipple press is different from that displayed byother chimpanzees in that he showed this behavior

Fig. 1. Puffy, massaging her left nipple with her right thumb.

Pan Africa News, 17(1), June 2010
frequently every day, and his pattern was meticulous.
Puffy’s nipple press was as meticulous as Alofu’s.
It is true that other chimpanzees also occasionallyshow nipple press: among adult females, Abi, Fatuma
and Wakusi; among adolescent males, Pim and Xmas;
among adolescent females, Qanat and Carmen (andRubicon, according to Takahisa Matsusaka); andamong adult males, Fanana. However, Puffy’s action
was so impressive that I hit upon the idea that Alofu
might have been Puffy’s father. By the way, Puffy’s
original closest associates, Pinky, Primus and Gwekulo,
never showed nipple press.
When I returned to Japan, I asked Eiji Inoue who
Puffy’s father was. Eiji had collected fecal samples
from as many individuals as possible and was able toclarify 11 pairs of father-offspring relationships via
DNA analysis7. Fortunately, Puffy was one of the few
youngsters whose father was clarified, and Eiji’s
answer confirmed my suspicion: he was Alofu! Puffy
had no close association with Alofu, and the possibilityof social learning from Alofu was small. Consequently,
the puzzle of the similarity between Puffy’s andAlofu’s patterns was solved.
There were mother-offspring pairs who sharednipple press habit, for example, Pim is Fatuma’s son,and Puffy’s father, Alofu, is Wakusi’s son. Althoughsocial learning was considered a possibility5
explaining the behavioral similarity, it may be thatgenetic influences should also be considered in thesecases. Thus, the nipple press behavior seems to be atleast partly genetically influenced.
On the other hand, Pim showed nipple press
when he was an adolescent but ceased doing this after
he became alpha male. Fanana did not show the
behavior when he was alpha male, but began to show
it by 2009, many years after he was dethroned in 2003.
Moreover, I have never seen any infants or juveniles
showing this behavior. Therefore, 9 dominance rank or
self-confidence and sexual maturity also seemed toinfluence the emergence of the nipple press.
There is another question: did other chimpanzees’nipple press patterns spontaneously occur, without
being influenced by Alofu’s frequent performances, orwere they influenced at least partially by Alofu?
Nipple press is a simple behavior that anyone canengage in without practice, so this is a difficult
question to address. When some behavior patterns
seemed to be in fashion in a group, we have too often
attributed it to social learning5. However, in the case ofpatterns giving physiological pleasure, we should
think twice whether or not it would be morereasonable to take genetic factors into primary11
consideration.
We should accumulate observational data ofnipple press behavior along with the association rates
of individuals as well as kinship analyses. Although an
indisputable answer may not be found, we could gaina better understanding of the influences of genetic
inheritance and social learning on behavioral patterns.
ACKNOWLEDGEMENTS
I thank Eiji Inoue for information from paternity
data; Kazuhiko Hosaka and Takahisa Matsusaka for
their sharing information on chimpanzees and their help
in the camp management during my stay at Kansyana
Camp in 2009 and their most helpful comments on thedraft of this paper; COSTECH, TANAPA and TAWIRI
for permission to do research at Mahale; and Mahale
Mountains National Park for logistic support. Field
research was supported by the JSPS Basic Scientific
Research A (# 19255008 to TN).
REFERENCES
1. 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.
2. Hosaka K. 1995. Mahale: A single flu epidemic killed
at least 11 chimps. Pan Afr News 2(2):3-4.
3. Nishida T. 1994. Review of recent findings on
Mahale chimpanzees: implications and future
research directions. In: Wrangham RW, McGrew WC,
de Waal FBM, Heltne PG (eds) Chimpanzee Cultures.
Harvard University Press, Cambridge, MA, pp.
373-396.
4. Marchant LF, McGrew WC. 1999. Innovative
behavior at Mahale: new data on nasal probe andnipple press. Pan Afr News 6:16-18.
5. Nishida T, Matsusaka T, McGrew W. 2009.
Emergence, propagation or disappearance of novel
behavioral patterns in the habituated chimpanzees ofMahale: a review. Primates 50:23-36.
6. Nishida T, Zamma K, Matsusaka T, Inaba A,
McGrew WC. 2010. Chimpanzee Behavior in the
Wild. An Audi-Visual Encyclopedia. Springer, Tokyo(in press).
7. Inoue E, Inoue-Murayama M, Vigilant L, Takenaka O,
Nishida T. 2008. Relatedness in wild chimpanzees:
the influence of paternity, male philopatry anddemographic factors. Am J Phys Anthropol
137:256-262.

Pan Africa News, 17(1), June 201012

T. Nishida, K. Zamma, T. Matsusaka, A. Inaba, W.C. McGrew

Chimpanzee Behavior in the Wild
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behavior patterns, from locomotion and posture to feeding,facial expression, gestures, grooming, and vocal
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First comprehensive list of behavior patterns ofchimpanzees with more than 1,000 entriesIncludes 700 video clips of chimpanzee behaviors on
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Color photos and drawings to supplement thedescriptions

Pan Africa News, Vol. 17, No. 1
Published in June, 2010
Address: Japan Monkey Centre
26 Kanrin
Inuyama, Aichi 484-0081
JAPAN
TEL: (+81)568-61-2327
FAX: (+81)568-62-6823
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ISSN: 1884-751X (Print), 1884-7528 (Online)