Pan Africa NewsThe Newsletter of the Committee for the Careand Conservation of Chimpanzees, and the Mahale
Wildlife Conservation Society
ISSN 1884-751X (print), 1884-7528 (online) mahale.main.jp/PAN/

JUNE 2012
P. A. N. EDITORIAL STAFF
Chief Editor:
Kazuhiko Hosaka, Kamakura Womenʼs University, Japan
Deputy Chief Editor:
Michio Nakamura, Kyoto University, Japan
Associate Editors:
Christophe Boesch, Max-Planck Institute, Germany
Jane Goodall, Jane Goodall Institute, USA
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
Takeshi Furuichi, Kyoto University, Japan
Editorial Secretaries:
Noriko Itoh, Kyoto University, Japan
Koichiro Zamma, Great Ape Research Institute, Hayashibara, Japan
Agumi Inaba, Mahale Mountains Chimpanzee Research
Project
Eiji Inoue, Kyoto University, Japan
Instructions for Authors:
Pan Africa News publishes articles, notes, reviews, forums,
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<ARTICLE>

Playing with His Leg: A Caseof Imaginary Social Play by an
Adolescent Male Chimpanzee at
Bossou?
Michio Nakamura
Kyoto University, Japan(E-mail: nakamura@wrc.kyoto-u.ac.jp)

VOL. 19, NO. 1

Contents<ARTICLE>
Playing with His Leg: A Case of Imaginary Social
Play by an Adolescent Male Chimpanzee at Bossou?
Michio Nakamura
1<NOTE>A Wild Chimpanzee Birth at Mahale
Koichiro Zamma, Tetsuya Sakamaki &Rashidi Shabani Kitopeni
3<NOTE>
Ecological Aspects of Chimpanzee Insectivory in the
Budongo Forest, Uganda
Sophie Hedges & William C. McGrew
6<NOTE>
Update on the Assirik Chimpanzee (Pan troglodytes
verus) Population in Niokolo Koba National Park,
SenegalJill D. Pruetz, Racine Ballahira, Waly
Camara, Stacy Lindshield, Joshua L.
Marshack, Anna Olson, Michel Sahdiako &Ulises Villalobos-Flores
8<NEWS>
Dr. Julius Keyyu Visits WRC
Gen’ichi Idani
11<NEWS>
Book Launch
12<BOOK INFO>
Long-Term Field Studies of Primates

12<BOOK INFO>The Ecological Impact of Long-Term Changes
in Africa's Rift Valley (Environmental Science,
Engineering and Technology)

13

INTRODUCTION
Human children sometimes create imaginary playmates. Together with other forms of pretense, imaginarysocial play is considered to be an essential part of humancognitive development. For example, it is argued that imaginary playmate fantasy occurs frequently among humanchildren and it facilitates social development by making
possible the practice of play frame negotiation1. Someauthors restrict imaginary social play to Homo sapiens,emphasizing that the complex interactions that characterize the play with an imaginary companion constitute1

2
a form of pretense that is beyond nonhuman animals2.
Besides “playmates,” imagination itself or pretense is an
important aspect of the human mind. However, this concept is rarely the main topic of the studies on nonhumanprimates (but see ref 3, 4). There are scattered reports oflanguage-trained or encultured apes showing imaginary
play or some form of pretense3,5–10. These observations
were often accompanied with the use of ASL or lexigrams
that could represent abstract concepts in the minds of theapes. In addition, when the imaginary play was “social”(i.e. oriented toward an imaginary companion), the directtargets of the behaviors were always human artifacts (suchas dolls) whose shapes often resemble, at least to observers, something animate.
There are also some anecdotal reports of imaginary
plays by the wild apes11–16 such that a juvenile chimpanzee
treated a log as if it were a baby. As some authors havepointed out, such examples have to be treated with caution. Great apes often handle objects for no particularpurpose, and some of the cases are not necessarily evidence of imaginary play. However, two Mahale cases15,16
are more convincing because the players exhibited playpanting although they were engaging in solitary play. This
vocalization is usually emitted when playing with a “social
companion”17 rather than during solitary play. In addition, some behavioral elements in these two cases (hitting,biting, mounting, aeroplane, etc.) are usually oriented toconspecifics. These two characteristics (play panting andsocially oriented behaviors) may be more reliable indicators for imaginary social play. Thus far, such examples
have been reported only from Mahale. In this paper, I
report the first case of possible imaginary social playfrom Bossou, Republic of Guinea that included these two
characteristics.

OBSERVATIONOn February 18, 2003, Tua (an old male) and Poni(a male born in 1993) were found together with otherchimpanzees at the Gban hill in the morning. At approximately 8:20 am, the two started to move separately awayfrom others to Gein hill across the road. At 10:55 am,
Poni walked backwards with his mouth open (showing
play face) to Tua. It is possible that Poni was soliciting
Tua to play; however, Tua did not respond, and Poni thengalloped in front of Tua. After feeding and moving for awhile, at 11:14 am, Tua began to sleep in the bush. Poni
continued to feed nearby. At 11:41 am, Poni’s play panting
was audible from the bush where he had been feeding. I
moved to a spot from where Poni was visible and started
to videotape his actions. After biting the pith of the undergrowth vegetation a little, he moved a few meters away
and resumed to play by himself by lying upward in thebush, holding his legs to his chest, and play panting repeatedly. At 11:45:46 am, he sat up, and with a play face,he pushed his own legs against the ground. Then after
putting his face close to the legs, he began to bite his left
leg (Figure 1a). He paused for a while and then resumed
biting his left leg for approximately 1 min. During this,his body was vibrating due to hard play panting.
At 11:47:14 am, he paused playing and did not display
a play face. After 10 s, he started observing his left leg,

Pan Africa News, 19(1), June 2012

Figure 1. Examples of socially oriented behaviors by Poni
during the play (all images drawn from the video
footage).
a) Biting his left ankle while supporting it with the righthand.
b) Moving the left foot back and forth toward theface while holding it with the left hand. It could beinterpreted that the leg was trying to escape while hewas trying to catch and bite it.
c) Hitting his left leg with the right knuckle repeatedly(3–12 times).
d) Pressing down the left leg against the ground withboth hands.
e) Raising the hips by pressing down the head in
his arms. A similar posture is also observed duringlocomotor play; however, in this context, it could
be interpreted that he was guarding himself fromsomeone.

and after extending the left hand to hold the leg and pull
it toward himself (Figure 1b), he again bit the leg for 15 s.
He sat straight and began to hit the left leg with the righthand (Figure 1c) and then he pressed the leg down against
the ground with both hands (Figure 1d). He then pirouetted and again pressed down the leg. He then hit the left
leg with the right hand in three intermittent sequences.
After stirring restlessly (presumably tickling himself) fora while, he got up and pressed his head down in his arms
against the ground while raising his hips (Figure 1e). At
11:51:20 am, upon hearing the calls of other individuals
some distance away, he screamed slightly and stopped
playing. He approached Tua and started to groom him.

DISCUSSION
It is notoriously difficult to define animal play18 because anything peculiar or anything different from thenormal behavioral categories could be described as play.
The above case is considered play due to the presenceof the play face and play panting, which are only seen inPan Africa News, 19(1), June 2012
the context of play. In addition, socially oriented behavioral elements such as biting, hitting, and pressing down
against the ground occurred frequently. Thus, this caseappears highly similar to what has been described as “imaginary social play” in Mahale chimpanzees14. However,since imagination cannot be directly observed, any proposed example of imaginary play can be criticized. For
example, Call and Tomasello19 were not convinced thecase of imaginary ant-dipping at Gombe11, because manyother interpretations are also possible. Similarly, what
may look like pretense can always be interpreted as something other than pretense20. Thus, alternative explanations
may be possible for Poni’s behavior. One possibility is that
Poni’s behavior was abnormal. However, Poni had nevershown any evidence of mental or physical illness. Anotherpossibility is that Poni’s play was just an extension of theusual solitary play (i.e. locomotor or object play). But thenwe need to explain why he delivered physical stimulation
to his own leg when there were many external objects that
could be played with. It is unlikely that he did this only
to play pant. Another difficulty with this interpretation isthat play panting does not occur reflexively in responseto physical stimuli. Self-tickling usually does not cause
humans to laugh spontaneously even when the stimulation is similar to that arising by being tickled by others. A
final possible interpretation is that this was truly a case ofimaginary social play, in which Poni treated his left leg as
if it were a social playmate, and directed socially oriented
behaviors toward it. Since we can never obtain direct
knowledge about others’ mental states, the imaginationof others (whether apes or humans) cannot be proved. We
cannot completely rule out alternative interpretations, but
nor can imagination be ruled out completely. This casemay be a rare case of imaginary play in wild great apes,
and the first one to be accompanied by play panting andsocially oriented behaviors in West African chimpanzees.
The unique aspect of this example is that the direct target
was the individual’s own body (the left leg, in particular)
instead of any external object. This indicates that Poni
played two different roles simultaneously—one was tobite and the other was to be bitten.
For Taylor and Carlson 2, imaginary companions (IC)are unique to humans because IC of human children areoften sustained for several months and usually have theirown personality. Thus far, we have no evidence that theimaginary playmates of chimpanzees are sustained orhave personalities; it may well be impossible to prove thepersonality of an imaginary playmate through observation. Thus, according to their criteria, chimpanzees would
not have IC even if they were capable of imagination.
However, Poni’s behavior can plausibly be interpreted as
imaginary social play, in that behavioral elements that areusually directed to live social playmates were repeatedly
directed to objects (including his own body and empty
space). It appears premature to exclude nonhuman animals from the realm of imaginary social play.

ACKNOWLEDGEMENTS
I thank DNRST of Guinea for permission to conduct thefield research at Bossou; G. Ohashi, G. Yamakoshi, and thestaff of IREB for their cooperation at the field; Y. Sugiyama and
T. Matsuzawa for giving me the opportunity to visit Bossou;

3
and T. Nishida for continuous support and guidance. The study
was financially supported by grants from Japanese MEXT(#12375003, #16255007 to T. Nishida, and #16770186 to M.
Nakamura) and by a grant for the biodiversity research of the
21st century COE (A14).

REFERENCES
1. Partington JT, Grant C 1984. Imaginary playmates and other useful fantasies. In: Play in Animals and Humans. Smith
PK (ed), Basil Blackwell, Oxford, pp. 217–240.
2. Taylor M, Carlson SM 2002. Imaginary companions andelaborate fantasy in childhood: Discontinuity with nonhuman animals. In: ref 4, pp. 167–180.
3. Jensvold MLA, Fouts RS 1993. Imaginary play in chimpanzees (Pan troglodytes). Hum Evol 8:217–227.
4. Mitchell RW (ed) 2002. Pretending and Imagination in
Animals and Children. Cambridge University Press,
Cambridge.
5. Hayes C 1951. The Ape in Our House. Harper, New York.
6. Fouts R, Mills ST 1997. Next of Kin. William Morrow, New
York.
7. Patterson F, Linden E 1981. The Education of Koko. Holt,
Rinehart & Winston, New York.
8. Savage-Rumbaugh S, Lewin R 1994. Kanzi: The Ape at the
Brink of the Human Mind. Wile, New York.
9. Matsuzawa T, Ueno A, Matsuno H, Hayashi M 2003.
Little chimpanzees (16): ‘Mimicry’ and ‘pretense.’ Kagaku
73:482–483 (in Japanese).
10. Miles HLW 1990. The cognitive foundations for reference
in a signing orangutan. In: “Language” and Intelligence inMonkeys and Apes. Parker ST, Gibson KR (eds), Cambridge
University Press, Cambridge, pp. 511–539.
11. Goodall J 1986. Chimpanzees of Gombe: Patterns of
Behavior. Harvard University Press, Cambridge, Mass., andLondon.
12. Wrangham R, Peterson D 1996. Demonic Males. Houghton
Mifflin, Boston.
13. Matsuzawa T 1997. The death of an infant chimpanzee at
Bossou, Guinea. Pan Afr News 4:4–6.
14. Hayaki H 1985. Social play of juvenile and adolescent chimpanzees in the Mahale Mountains National Park, Tanzania.
Primates 26:343–360.
15. Byrne R 1995. The Thinking Ape. Oxford University Press,
Oxford.
16. Nishida T, Kano T, Goodall J, McGrew WC, Nakamura M
1999. Ethogram and ethnography of Mahale chimpanzees.
Anthropol Sci 107:141–188.
17. van Lawick-Goodall J 1968. A preliminary report on expressive movements and communication in the Gombe
Stream chimpanzees. In: Primates. Jay PC (ed), Holt,
Rinehart & Winston, New York, pp. 313–374.
18. Bekoff M, Allen C 1998. Intentional communication andsocial play: How and why animals negotiate and agree toplay. In: Animal Play. Bekoff M, Byers JA (eds), Cambridge
University Press, Cambridge, pp. 97–114.
19. Call J, Tomasello M 1996. The effect of humans on the
cognitive development of apes. In: Reaching into Thought.
Russon AE, Bard K A, Parker ST (eds), Cambridge
University Press, Cambridge, pp. 371–403.
20. Gomez JM, Martin-Andrade B 2002. Possible precursorsof pretend play in nonpretend actions of captive gorillas(Gorilla gorilla). In: ref 4, pp. 255–268.

<NOTE>A Wild Chimpanzee Birth at
MahaleKoichiro Zamma1,3, Tetsuya
Sakamaki2,3 & Rashidi Shabani

Pan Africa News, 19(1), June 2012

4

Kitopeni3
1. Great Ape Research Institute, Hayashibara, Japan
2. Primate Research Institute, Japan
3. Mahale Mounatins Chimpanzee Research Project(E-mail: zamma@gari.jp)

INTRODUCTION
Wild chimpanzee births are difficult to observe because females tend to hide during the birthing process to
avoid others1, 2. On 26 May 2000, Juno, a female chimpanzee at Mahale Mountains National Park, Tanzania, gavebirth. Although we did not see the exact moment of birth,
we report our observations of her behavior immediately
before and after the birth.

OBSERVATIONSBefore birth
At 08:14 on 26 May 2000, we began to observe thechimpanzees of M group in Mahale. At 08:21, one researcher (KZ) started to follow an adult male called
Kalunde. Around 08:30, Juno appeared uttering pantgrunts to Kalunde. She then climbed a tree and lay on a
day bed made of one branch. The chimpanzees moved
north and ate the fruit of Psydrax parviflora or Uvaria
angolensis.
At 11:46, Juno reappeared. She crouched on the
ground in front of Kalunde, and he began to groom her.
After a few seconds, Alofu, an adult male, approachedand peered at Juno’s genitals. Kalunde also tried to peerat them, but she turned around once to prevent this. Her
genital skin was not swollen, seemed dry, and showedno sign that her water had broken. She had not given
birth at that time. When Juno lay on her left side, Alofu
and Kalunde examined her genitals for 13 seconds.
Subsequently, she lay on her back, and the males started
grooming her. She did not reciprocate the grooming. At
11:47, Juno rolled over and lay on her left side, and thenagain lay on her back at 11:48. Alofu changed his position
to follow her genitals. At 11:49, Juno lay on her left sideagain, and at 11:51 she turned around and crouched. Alofuchanged his position again, following her genitals, and he
and Kalunde continued to groom her. At 11:57, Kalunde
stopped grooming and lay in the same location, but Alofu

Figure 1. Juno (left) and her newborn. Totzy (right)
watched them.

continued to groom Juno. At 12:01, Kalunde groomed
Alofu for a short time and then left. When KZ left, following Kalunde, Juno was still being groomed by Alofu.
After birth
When a researcher (RK) observed Juno in the tree at
14:44, she was already nestling her newborn in her arms.
RK had followed his focal animal, Masudi (an adult male),to the location and he also found Alofu on the ground.
Juno climbed down from the tree, uttered pant-grunts to
Alofu, and climbed another tree. Then, Fanana (the alphamale) came to the tree, displayed, and charged Juno. She
screamed and climbed higher. After Fanana left, Juno
made a bed and started to rest.
At 15:03, researchers KZ and TS arrived at the tree
while following their focal animals. TS had been following an adult male chimpanzee named Bonobo since thatmorning. Juno appeared from the bed with her newborn
and touched her genitals five times while sat on a branch
20 m above the ground. KZ took video on it using SONY
DCR-PC7. Four adult females (Caliope, Totzy, Nkombo,
and Zola), an adult male (Carter), and four immature
chimpanzees (Primus, Ivana, Jidda, and Christmas)
watched Juno in the tree. Caliope was a multiparous female with a 2-year-old offspring, and Totzy, Nkombo,
and Zola were nonparous females with swollen genitals.
Jidda was the 5-year-old offspring of Juno. At 15:05,
Juno put the end of the umbilical cord in her mouth andpulled on it as if she was trying to the suck fluid insideit (Figure 1, Video 1: available online at mahale.main.jp/
PAN/19_1/19(1)_02.html). The umbilical cord was about
40 cm long and the other end was connected to her newborn. The placenta was not observed. Then, Juno moved
on woody vines while holding the umbilical cord in her
mouth (Figure 2). The newborn was clinging to her abdomen, held there with Juno’s thigh and hand. Primus stared
at Juno from a distance of 3 m. At 15:07, Juno defecatedand urinated, then lay on her back on a woody vine. When
Caliope approached Juno while fleeing from Primus, Junouttered soft barks and moved on the tree while holding theumbilical cord in her mouth. At 15:09, Primus and Carter
started to display in the tree, but they hastily climbed
down and uttered pant-grunts to Fanana, who had ap-

Figure 2. Juno and the umbilical cord.

Pan Africa News, 19(1), June 2012
peared beneath the tree. Carter also uttered pant-screams
to Fanana, and then Juno and another chimpanzee utteredpant-hoots. At 15:12, Fanana left the location and someof the other chimpanzees also started to leave the tree.
At 15:21, Caliope also left. Juno uttered “hu, hu” and thenewborn uttered “fa, ha-ha.” The newborn was female.
Terrestrial movement
At 15:23, Juno descended the tree and walked on the
ground with her newborn, which she held with her righthand. A researcher (TS) followed her. At 15:25, Juno
shifted to holding her newborn with her left hand andwalked tripedally while trailing the umbilical cord. She
passed in front of Masudi uttering pant-grunts, and saton the ground. Masudi approached Juno from behind.
She showed a grin face while uttering soft pant-grunts.
She looked frightened. Masudi stood quadrupedally andpeered at her from a distance of 10 cm. At this time, Pimu(an adolescent male) approached and sat in front of Juno.
When Juno left, Pimu attacked her back. Juno ran awayuttering screams and barks. Pimu left, while displaying.
Caliope came to Juno’s side, like a guard, uttering vocalizations. Juno and Caliope walked together in a different
direction from the others.
At 15:30, Caliope had left and Juno walked alone. She
ate U. angolensis fruit. She pulled the umbilical cord withher right hand. At 15:42, she lay on her back. The newborn sucked Juno’s left nipple. When the newborn almost
slid off, Juno held her up. The newborn uttered “hu, hu.”
At 15:44, Juno sat up. TS heard the vocalizations of otherchimpanzees some distance away. At 15:52, Juno pickedup the umbilical cord and the newborn uttered “hu, hu.”
At 15:53, Juno put the umbilical cord in her mouth andchewed on it. At 15:57, Juno started to walk with the newborn, trailing the end of the umbilical cord. At 15:58, sheentered a thick bush and TS stopped following her.
We observed 50/51 individuals in M group that day.
Examination of the bed
At 15:56, researcher KZ climbed up to observed theempty bed. It was 80 cm in diameter and the placenta was
not found on it.

DISCUSSION
Juno gave birth between 12:01 and 14:44 on 26 May
2000 while moving with almost all members of her unit
Table 1. The partners and number of copulations with
Juno observed from November to December
2000 (TS unpublished data).

＿＿＿＿＿＿＿＿＿
Alofu
1
Bonobo 4
Carter 2
Dogura 1
Fanana 4
Hambi 5
Kalunde 2
Masudi 2
Pimu
3
Primus 3＿＿＿＿＿＿＿＿＿

5group. Wild chimpanzee births are observed extremely
rarely1–3. Juno immigrated in 1987 and had given birth tofour offspring (MMCRP unpublished data). She had spenta long time in M group and had given birth many times,
so she might have had the confidence to give birth thefifth time in broad daylight.
The umbilical cord was observed, but the placenta
was not found. Previous reports of chimpanzee births
have described the delivery of the placenta 13–18 minutes
after birth, and the mother’s consumption of it beginning
0–4 minutes later1,3. Juno must have eaten the placenta
shortly after she delivered.
Based on a 229-day gestation period 4, Alofu was
the likely father of Juno’s offspring because only Junoand Alofu were not observed from 26 September to 11
October 2000 and were thought to have formed a consortship. However, Juno was observed to copulate at least 27
times with 10 males subsequently (Table 1. TS unpublished data), so the paternity was confused among thesemales. Because the males had the idea that they were thefather of her newborn, they may not have delivered terrible attack on Juno and her newborn though some of themdisplayed near Juno and charged her.
It is interesting that Alofu, the probable father, spenttime peering at Juno’s genitals before the birth, although
they were not swollen and lacked any visible sign of parturition. The urinary estrone conjugate (E1C) of female
chimpanzees peaks during genital swelling and just before birth, whereas urinary pregnandiol glucuronide (PdG)
remains at low levels during swelling and increases justbefore parturition5. Alofu may have smelled an odor indicating swollen genitals or imminent parturition.

ACKNOWLEDGEMENTS
We thank TAWIRI, TANAPA and COSTECH for permission of our fieldworks, Rashidi H, Mtunda M, Hamisi B, andother member of MMCRP for their help in the field, and Fujita
S, Nakamura M, Hosaka K and Itoh N for their valuable comments. This study was financially supported by grants from the
COE program of Monbusho (#10CE2005 to O. Takenaka) andthe Leakey Foundation (to T. Nishida).

REFERENCES
1. Goodall J, Athumani J 1980. An observed birthe in a freeliving chimpanzee (Pan troglodytes schweinfurthii) inGombe National Park, Tanzania. Primates 21: 545–549.
2. Reynolds V 2005. The Chimpanzees of the Budongo Forest:
Ecology, Behaviour, and Conservation. Oxford, New York.
3. Kiwede ZT 2000. A live birth by a primiparous female
chimpanzee at the Budongo forest. Pan Africa News
7:23–25.
4. Martin DE, Graham CE, Gould KG 1978. Successful artificial insemination in the chimpanzee. Symp Zool Soc Lond
43:249–260.
5. Shimizu K, Douke C, Fujita S, Matsuzawa T, Tomonaga M,
Tanaka M, Matsubayashi K, Hayashi M 2003. Urinary steroids, FSH and CG measurements for monitoring the ovarian cycle and pregnancy in the chimpanzee. J Med Primatol
32:15–22.

Pan Africa News, 19(1), June 2012

6

<NOTE>

Ecological Aspects of Chimpanzee
Insectivory in the Budongo
Forest, Uganda
Sophie Hedges & William C.
McGrew
Department of Archaeology & Anthropology, University ofCambridge, UK(E-mail: wcm21@cam.ac.uk)

INTRODUCTION

All long-term studies of chimpanzees (Pan troglodytes) show that they habitually eat insects, most oftensocial insects obtained by extractive foraging with simple tools1. This generalisation holds across Africa from
Senegal to Tanzania, but Ugandan populations are exceptional: Except for Kalinzu with ant dipping for Dorylus2,
no other Ugandan long-ter m study site (Budongo,
Kanyawara, Ngogo, Semliki) shows regular insectivory,
technically-aided or otherwise3. At Budongo, only occasional cases of eating Cubitermes have been recorded4.
This dearth is puzzling, especially when these sites arecompared with those (Gombe, Mahale) of neighbouring
Tanzania, which have well-documented and varied elementary technology5,6.
Lack of insectivory could reflect basic environmental
constraints: Absence or scarcity of prey species, or absence or scarcity of raw materials for tools7. Less likely
alternative explanations are differences between Ugandan
and non-Ugandan populations in terms of genome, diet,manual dexterity, intelligence, or appetite. The presenceof ant dipping at Kalinzu casts doubt on all these alternatives. Finally, such inter-population differences could
result from differences in cultural knowledge, as has beenfound elsewhere8. That is, some populations of chimpanzees may not have discovered that some insects are edibleor can be got with technical assistance.
Our study sought to explore the role of environmental
constraints in the absence of regular insectivory in the
Budongo Forest Reserve, Uganda.

METHODS
The study was done in the Sonso region Budongo
from 14 July–8 September, 2011. Subjects were 80+ wellhabituated chimpanzees (Pan troglodytes schweinfurthii)of the Sonso community9. SH did 12 line-transects of 6
m width, each of 500 m length, along the existing grid
system of trails; six ran north-south and six ran east-west.
Sites of transects were balanced for the two main forest
types, mixed (4,260 m) and swamp (1,740 m). Distanceswere measured by hip-chain, while another person
scanned for evidence of social insects. Total area surveyed
was 3.6 ha.
The following taxa (all k now n to be eaten bychimpanzees elsewhere in East Africa) were sought:
Cubitermes, Macrotermes, and Pseudacanthotermes
termites; Dorylus (army) and Oecophylla (weaver) ants;
and Apis (honey) bees. SH checked termite mounds for
activity and measured their height and circumference;

this allowed calculation of their volume, either as a cylinder or cone. Encounters with Dorylus ants on trails werenoted by GPS data on time and place. Mounds and nestsof all species were monitored weekly, after initial datacollection and surface clearing of debris. On each visit,
SH checked for insect activity and signs of chimpanzee orother predator presence or exploitation. Specimens of preyspecies were preserved and identified later by specialists.
SH surveyed raw materials suitable for extractive
foraging probes, that is, woody and non-woody vegetation
with straight, elongate dimensions. Data-collection entailed marking a circle of 5 m radius around the resource
and counting all plants presenting potential raw materials,
in three categories: woody tree or shrub, vine, or monocotyledon (e.g. grass, sedge)10.
To measure the chimpanzees’ consumption of insects,
both direct and indirect data were collected. SH logged 80
hr of direct observation of opportunistically encounteredparties of chimpanzees, using continuous recording of
feeding or interacting with insects11. For indirect data, SH
collected all fresh faecal samples (n = 26) from chimpanzees encountered; these were sealed in ziplock bags andlater sluiced in running water to detect undigested foodfragments12.
To estimate productivity, that is, amount of termites
available to harvest, SH randomly sampled 10 Cubitermes
and two Pseudacanthotermes mounds, removing about
150 cm3 from their tops. Once sealed in a ziplock bag, thesoil was broken into chunks of less than 1 cm 3; all termites, by caste, were counted in the contents7. Also, SH
made and used flexible probes modelled on chimpanzee
fishing probes, to fish termites from these mounds. To
measure the payoff from termite fishing, efficiency, success rate, and error rate were calculated12.
Because sample sizes were small and data non-normally distributed, all statistical tests were non-parametric.
Alpha was set at 0.05, and all tests were two-tailed.
For specific details of methods, see Hedges13.

RESULTS

Cubitermes ugandensis mounds were found at amean density of 34.4 mounds/ha (SEM = 8.17). Neither
Macrotermes bellicosus nor Pseudacanthotermes spiniger mounds were found on transects, but two mounds ofeach genus were found and monitored elsewhere. Twochimpanzees were seen to eat soil from mounds, one of
Pseudacanthotermes and one of Cubitermes. Two mounds(one of each genus) showed damage from chimpanzee
predation, but the two Macrotermes mounds showed nosigns of chimpanzee use.
Volu me of ter mite mou nds var ied g reatly but
did not differ across forest types. Mean volume of
Macrotermes mounds was almost 300 times that of
the average Cubitermes mound, and mean volume of
Pseudacanthotermes mounds was almost seven times as
big.
Encounter rate over 6 days of Dorylus (wilverthi or
kohli) columns on trails was 0.20 columns per km and 0.42
columns per hr. Neither Apis mellifera nor Dorylus nests
were found on transects, but one of each was found andmonitored elsewhere.

Pan Africa News, 19(1), June 2012
Raw mater ials for f ishing probes were superabundant. Cubitermes mounds afforded a mean of 580
tool sources per mound, while the numbers of sources
for other insect prey species were: Macrotermes (155),
Pseudacanthotermes (460), Dorylus (440), and Apis (684).
Availability of raw materials did not differ across preyspecies, but overall, woody plants predominated overmonocots or vines in abundance.
None of the 26 faecal specimens yielded insect
remains.
The 12 mounds assessed for termite availability yielded an average of 104.8 termites per sample, but the rangeof values was huge (0 –361).
SH’s fishing from Cubitermes mounds was minimally productive, as the passageways were narrow
and the soldiers passive. Fishing was a far less useful
harvesting technique than detaching portions of themound, with mean yields of 2.4 versus 123.5 termites.
Pseudacanthotermes showed the reverse: Fishing was far
more productive than detaching soil, with mean yields of
139 versus 11.5 termites. Fishing the two Macrotermes
mound yielded almost nothing (mean of 2.5 termites).

DISCUSSION
Availability of prey or of raw materials seems
not to be an obstacle to Budongo chimpanzees using tools to obtain insects. The low abundance ofmounds of Macrotermes and Pseudacanthotermes iswithin the range of densities at other sites where thesegenera are f ished: Bilenge (0.35/ha, Macrotermes;
1.03, Pseudacanthotermes); Campo (0.68/--); Gombe(0.40/2.04), etc. Similarly, the density of Cubitermes
mounds at Budongo, at 34.4 mounds/ha, where the termites are eaten without tools, is intermediate in a wide
range of values: 0.8/ha at Gombe at one extreme, to 233.8/
ha at Lui Kotale at the other.
Similarly, encounter rates for Dorylus at Budongo,measured by either distance or time, are comparable tothose from other sites. For distance, Budongo’s encounterrate of 0.2/km is close to Gashaka’s dry season rate of
0.21, where the highest rate of consumption of Dorylus bychimpanzees has been recorded. For time, Budongo’s rateof 0.42/hr also resembles Gashaka’s at 0.4314.
Too few data are available to compare abundance ofApis nests between Budongo and other sites. Similarly,
too few data have been published or were obtained here toallow confident comparison of termite mound volumes.
Consumption of termite soil (and termites) by chimpanzees varies hugely. Budongo’s rate of 0.79/100 hr ofsuch geophagy is higher than that at Kibale (0.52) but
much lower than that at Kasoje (4.07)15. This report ofSonso chimpanzees eating the soil of Pseudacanthotermes
appears to be a first for this genus in Uganda.
Raw materials of all types for extractive probes
are readily available. To take the most pertinent case,
Budongo’s Macrotermes mounds afforded 155 raw material sources on average, compared with a range from 37 at
Assirik to 228 at Semliki.
Absence of termites in chimpanzee faecal samples
replicates the results of an earlier study4, confirming that
insectivory is rare.

7
Technical (i.e. fishing) versus non-technical (i.e.
detachment) acquisition of termites seems not to havebeen compared systematically before this study. The closest result to ours seems to be that tools and techniques
used to obtain Dorylus differ according to species ofant8. But until the chimpanzees of Budongo learn to eat
Pseudacanthotermes and Macrotermes, these differences
remain hypothetical.
In conclusion, the most likely explanation for the lackof extractive foraging for insects by Budongo chimpanzees is lack of cultural knowledge.

ACKNOWLEDGEMENTS
We thank: Uganda Wildlife Authority (UWA) and Uganda
National Council for Science and Technology (UNCST) for
permission to do research; K. Zuberbuehler and G. Muhanguzi
for permission to work at BCFS; Corpus Christi College;
Department of Biological Anthropology; and the Mosley, Frere,and Worts Funds of University of Cambridge for funding; R.
Ogen, G. Muhumuza, C. Hobaiter, and B. Fallon for field assistance; C. Schöning and P. Nyeko for insect identifications.

REFERENCES

1. McGrew WC 1992. Chimpanzee Material Culture:
Implications for Human Evolution. Cambridge University
Press, Cambridge.
2. Hashimoto C, Furuichi T, Tashiro Y 2000. Ant dipping andmeat eating by wild chimpanzees in the Kalinzu Forest,
Uganda. Primates 41:103–108.
3. Watts DP 2007. Tool use by chimpanzees at Ngogo, Kibale
National Park, Uganda. Int J Primatol 29:83–94.
4. Newton-Fisher NE 1999. The diet of chimpanzees in the
Budongo Forest Reserve, Uganda. Afr J Ecol 37:344–354.
5. Goodall J 1986. The Chimpanzees of Gombe: Patterns of
Behavior. Harvard University Press, Cambridge, Mass., andLondon.
6. Nishida T 2012. Chimpanzees of the Lakeshore: Natural
History and Culture at Mahale. Cambridge University
Press, Cambridge.
7. McGrew WC, Marchant LF , Beuerlein MM, Vrancken
D, Fruth B, Hohmann G 2007. Prospects for bonobo insectivory: Lui Kotal, Democratic Republic of Congo. Int J
Primatol 28:1237–1252.
8. Schöning C, Humle T, Möbius Y, McGrew WC 2008. The
nature of culture: Technological variation in chimpanzee
predation on army ants revisited. J Hum Evol 55:48–59.
9. Reynolds V 2005. The Chimpanzees of Budongo Forest:
Ecology, Behaviour and Conservation. Oxford University
Press, Oxford.
10. Martin P, Bateson P 2007. Measuring Behaviour: An
Introductory Guide. 3rd edition. Cambridge University
Press, Cambridge.
11. McGrew WC, Marchant LF, Phillips CA 2009. Standardised
protocol for primate faecal analysis. Primates 50:363–366.
12. McGrew WC, Marchant LF 1999. Laterality of hand use
pays off in foraging success for wild chimpanzees. Primates
40:509–513.
13. Hedges S 2012. Ecological aspects of chimpanzee insectivory in the Budongo Forest, Uganda. Final-year dissertation, University of Cambridge.
14. Fowler A, Sommer V 2007. Subsistence technology of
Nigerian chimpanzees. Int J Primatol 28:997–1023.
15. Mahaney WC, Milner MW, Sanmugadas K, Hancock
RGV, Aufreiter S, Wrangham R, Pier HW 1997. Analysis
of geophagy soils in Kibale Forest, Uganda. Primates
38:159–176.

Pan Africa News, 19(1), June 2012

8

<NOTE>

Update on the Assirik
Chimpanzee (Pan troglodytes
verus) Population in Niokolo
Koba National Park, Senegal
Jill D. Pruetz1,2, Racine Ballahira3,
Waly Camara2, Stacy Lindshield1,4,
Joshua L. Marshack5, Anna
Olson1, Michel Sahdiako2 & Ulises
Villalobos-Flores2
1 Department Anthropology, Iowa State University, Ames,USA
2 Fongoli Savanna Chimpanzee Project, Kedougou, Senegal
3 Departmentof Research and Management, Niokolo Koba
National Park, Tambacounda, Senegal
4 Ecology and Evolutionary Biology Program, Iowa State
University, Ames, USA
5 Department of Anthropology, Washington University, St.
Louis, USA(E-mail: pruetz@iastate.edu)

Figure 1. Location of Niokolo-Koba National Parkin Senegal.

INTRODUCTION
The Assirik, Senegal chimpanzee population wasstudied extensively by McGrew and colleagues in the
Stirling African Primate Project (SAPP) in the late 1970s1
and surveyed by the Miami Assirik Pan Project (MAPP)
in 20002. These chimpanzees (thought to represent a single community3), within the Niokolo Koba National Park(or Parc National du Niokolo Koba, PNNK), remain the
only protected population in the country, with most of
Senegal’s chimpanzees living in unprotected areas4. In
2012, the Iowa State Assirik Primate Project (ISAPP) surveyed Assirik and outlying areas in the PNNK to assess
chimpanzee density and to explore the possibility of establishing a research program there in collaboration with
the National Parks Department of Senegal and Universityof Cheikh Anta Diop in Dakar. One objective in this study
was to re-survey the Assirik area in order to compare ourresults to data collected in 2000.

METHODS
Niokolo Koba National Park is in southeaster n
Senegal (12o53’ N, 12o44’ W) (Figure 1). The habitat is
Sudano-Guinean mixed woodland and savanna, characterized by an extensive 7-month dry season and less than
1,000 mm of rainfall per year. The park has four diurnal
non-human primate species (see Table 2) and two nocturnal species (Galago senegalensis, Potto perodicticus).
Several potential mammalian predators on chimpanzees
exist in the PNNK, including lion (Panthera leo), leopard(Panthera pardus), spotted hyena (Crocuta crocuta) andwild dog (Lycaon pictus)5.
We combined systematic line transect and reconnaissance sampling of chimpanzee nests in an effort to replicate methods from the MAPP study2. However, with lessthan half of the time available to conduct the survey, we
did only two rather than four 2-km transects that radiatedfrom the summit of Assirik and did not repeatedly sample
areas. On 12 different days, from January 26 –February
2, 2012, we surveyed 115.7 km (78.7 km on foot, 37 km

by vehicle) during daylight hours for chimpanzee nests.
We sampled the Mt. Assirik area as well as areas to thesouth and north of this summit (Table 1, Figure 2). This
included two 2-km transects at the Assirik summit and6.7 km in two forested valleys (Stella’s and Lion Valleys),in addition to searching wooded areas. In order to provide
a measure of potential chimpanzee predators, prey andcompetitors, we calculated encounter rates (number ofsightings or individuals per km) with mammals (excluding
rodents and bats), on foot (102.4 km) and from the vehicle(448.6 km) between the Ranger Post to the southeasternsection of PNNK. Vehicle surveys could cover the same
area more than once and included encounters with animals seen on the Tambacounda-Kedougou highway running through the PNNK.
We made several assumptions in our analyses of nest
densities. We assumed all nests were less than 120 days
old and that each weaned chimpanzee built a new nest
each night. We used a formula revised from Marchesi andco-workers6: Density = (# nests/area surveyed) (1/mean
nest duration [120 days]) to calculate the density of nestbuilding chimpanzees. We weighted our values according
to habitat type since the Assirik summit transects bisected
only woodland and grassland habitats, and most nests
were found in gallery forests. Therefore, we calculated
nest density in woodland (including bamboo woodland =42% of area) and gallery forest (3% of area) separately inorder to produce a more accurate measure of chimpanzee
density in this savanna mosaic. Percentage habitat values
are from Baldwin and co-workers7. No nests were foundin grassland habitats during the transect surveys, which
accounts for 55% of the area used by Assirik chimpanzees7. We used data from the 2.77 km transect throughLion Valley to represent chimpanzee nest density in gallery forests and used the two 2-km transects from thesummit of Assirik to represent woodland habitats, multiplying these values by 3% and 42%, respectively. This
procedure replicates MAPP methods2.

Pan Africa News, 19(1), June 2012

9

RESULTSWe recorded 840 nests built bychimpanzees in the PNNK (Table 1,
Figure 2). Additionally, we heard atleast one chimpanzee pant-hooting
during a transect survey on the eastern slope of Assirik. Chimpanzee
nest density in the PNNK was estimated at 1.28 individuals per km2.
We recorded 16 mammal species during 106 sightings (0.19 perkm) with 476 individuals (0.86 perkm) (Table 2). We also found recent
traces of poachers (e.g. shotgun
shells, footprints, sounds of gunfire).
During one foot survey, we encountered a group of poachers and then
immediately left the area for safety
reasons.

DISCUSSION
Although our results from thisbrief survey indicate that the chimpanzee population in the PN NK
increased since the MAPP survey,
from 0.13 nest-building chimpanzees
per km 2 calculated in 20002 to 1.28
in this study, such an interpretation
should be made with caution. The
results reported here accounted forless than 7 km of line transects, compared to the 13.7 km surveyed in the2000 study2. Plumptre8 recommend
surveying at least 200 km for forestdwelling primates. However, effective sampling distances in a savanna
environment are likely to be shorter
because detection distances (i.e.
strip width) are larger. In the MAPP
survey2, a detection distance of 36 m
was calculated based on nest sighting
distances, suggesting a strip width of

Figure 2. Locations of nest survey areas and number of nests
encountered within Niokolo-Koba National Park.

Table 1. Surveys of chimpanzee nests and mammals in Niokolo Koba National Park. Nest survey
distances are in parentheses.

Date
16 Jan17 Jan18 Jan19 Jan23 Jan24 Jan25 Jan26 Jan30 Jan31 Jan
1 Feb
2 Feb
Total

Km surveyed Km surveyed Nest countArea surveyed
by vehicle
on foot
79(26)4(4)0Southeast of Assirik
0(0)23(23)
49
Southeast of Assirik
35(0)10(5.3)0Southeast of Assirik
44(0)
0(0)
0
Highway69(5)3.8(3.8)
78
Assirik woodlands
0(0)9.5(9.5)
75
Assirik area, Stella's Valley
0(0)11.4(11.4)
213
Mount Assirik area
69(0)1.7(1.7)
121
Assirik gallery forest and woodland
69(0)8.3(4.2)
30
Assirik gallery forest and woodland
6(6)12.4(7.6)
126
Mount Assirik area
8.6(0)
14(5.7)
68
Assirik area, Lion Valley
69(0)4.3(2.5)80
Tower highway gallery forest and woodland
448.6(37)102.4(78.7)
840

GPS point
12°49.373 N, 12°39.782 W
12°51.171 N, 12°42.945 W
12°49.400 N, 12°39.818 W
13°04.511 N, 12°43.384 W
12°53.738 N, 12°42.360 W
12°53.012 N, 12°43.670 W
12°52.682 N, 12°45.130 W
12°53.532 N, 12°42.589 W
12°57.469 N, 12°43.562 W
12°52.401 N, 12°46.156 W
12°51.395 N, 12°46.864 W
13°02.407 N, 12°40.278 W

10

Pan Africa News, 19(1), June 2012

72 m provided a reliable estimate of Table 2. Mammal species recorded in Niokolo Koba National Park*.
chimpanzee nest density in this envi# times# individualsronment. Nonetheless, given the brief Species
encounteredencounterednature of our study, our comparative
Baboon (Papio hamadryas papio)
27
248
analysis should be used primarily as a
Vervet monkey (Chlorocebus aethiops)
26
108
relative measure of the presence andPatas monkey (Erythrocebus patas)
726
location of chimpanzees in the PNNK
Warthog(Phacochoerus
africanus)
7
26
over the years. However, we found16
23
a large number of chimpanzee nests Oribi (Oribi oribi)
Derby
Eland
(Taurotragus
derbianus)
217during a relatively brief survey, which
Duiker species (Cephalophus spp.)
912
is encouraging.
1
4
We recorded more nests in 2012 Banded mongoose (Mungos mungo)
3
3
relative to the 2000 survey, although Bushbuck (Tragelaphus scriptus)
nest encounter rates (nests per km) Hartebeest (Alcelaphus buselaphus)
12were higher in 2000. A total of 29 km Grimm’s duiker (Sylvicapra grimmia)
1
1(on foot) and 54 km (via vehicle) were Roan antelope (Hippotragus equinus)
1
1
surveyed during the MAPP study 2 Slender mongoose (Galerella sanguinea)
11compared to the current effort of 78.7 Egyptian mongoose (Herpestes ichneumon)
1
1
km and 37 km surveyed by foot and Mongoose (sp. indet.)
1
1vehicle, respectively. We recorded Civet? (Viverra civetta?)
1
1
840 nests along 115.7 km (7.3 nests
Golden cat? (Caracal aurata?)
1
1
per km surveyed), while MAPP reTotals
106476
corded 736 nests along 83 total km (8.9*Species heard but not seen were leopard (Panthera pardus),nests per km surveyed). We attributespotted hyena (Crocuta crocuta) and chimpanzee (Pan troglodytes verus)
the lower encounter rate in 2012 to reconnaissance sampling more than 10
km southeast of the Assirik summit, where no nests were uted to rising birth rates and immigration from surrounding countries. Large-scale mining is increasing here, andobserved (Figure 2).
Based on our 2012 records of nests in new areas sur- smaller-scale artisanal mining attracts people from otherveyed relative to the 2000 study, we suggest that at least countries who may not have the same cultural values that
two chimpanzee communities use the PNNK, in contrast have resulted in the protection of Senegal’s chimpan4
to the findings of Tutin and co-workers3. We recorded zees via taboos against eating them . The large mammal
nests more than 20 km north of Mount Assirik (Figure 2). population in the PNNK is thought to be under increased
At the Fongoli site, less than 60 km from Assirik (Figure pressure from poachers, and their presence could also af1), the chimpanzees’ home range is over 86km 2, and the fect the behavior of chimpanzees in this area, althoughfurthest distance between points used by this community they are not yet thought to be the target. We found moreis around 10 km (Pruetz & Wessling, in prep.). We hy- evidence of poachers in our 12 days in the PNNK thanpothesize that PNNK chimpanzees exhibit similar rang- JDP observed in over 2 months during the 2000 study.
ing patterns, and may in fact have smaller home ranges As mining continues to develop in Senegal, the market
than chimpanzees outside of the PNNK since they do not for bushmeat is likely to increase even more, indicating
experience anthropogenic disturbance, which accounts for the need to better protect this important area of wildlife
5% of the Fongoli range9. Although both the 2000 and the diversity.
2012 surveys occurred during the dry season (February–
March and January, respectively), data on Fongoli chim- ACKNOWLEDGMENTS
We thank the Republic of Senegal, Direction des Parcs
panzees indicate that savanna chimpanzees in Senegal
Nationaux,
especially Docteur Djibril Diouck, Commandant
use their home ranges seasonally and cyclically. Fongoli Mamadou Sidibe,and the rangers at Niokolo Koba Post for aschimpanzees range most widely during the early dry sea- sistance. We thank Gray Tappan for information on the locationson, when baobab is fruiting. It is possible that Assirik of chimpanzees in the PNNK, Erik Otarola-Castillo for technichimpanzees do so as well, ranging more widely from the cal advice, and WCM McGrew for helpful comments on thissummit during the early dry season (November–January) manuscript.
compared to the late dry season (February–April) due
to the widespread dispersion of baobab trees, a key food REFERENCES
1. McGrew WC, Baldwin PJ, Tutin CEG 1981. Chimpanzees
source for Fongoli chimpanzees10. In any case, the harsh
in a hot, dry, and open habitat: Mt. Assirik, Senegal, Westsavanna environment coupled with confirmed poaching
Africa. J Hum Evol 10:227–244.
activity within the park’s borders underscores the need 2. Pruetz JD, Marchant LF, Arno J, McGrew WC 2002.
Survey of savanna chimpanzees (Pan troglodytes verus) infor increasing conservation and management efforts forSoutheastern Senegal. Am J Primatol 58:35–43.
Assirik chimpanzees. Given the increasing human popu- 3. Tutin CEG, McGrew WC, Baldwin PJ 1983. Social organilation in Senegal, it is possible that the human community
zation of savanna-dwelling chimpanzees, Pan troglodytes
verus, at Mt. Assirik, Senegal. Primates 24:154–173.
surrounding PNNK will exert increasing competitive and
4. Carter J, Ndiaye S, Pruetz J, McGrew W 2003. Senegal.
predation pressures on chimpanzees in the years to come.
In: West African Chimpanzees: Status Sur vey and
The increasing human population of Senegal is attribConservation Action Plan. Kormos R, Boesch C, Bakarr

Pan Africa News, 19(1), June 2012
MI, Butynski TM (eds), IUCN, pp. 31–39.
5. Di Silvestre I, Novelli O, Bogliani G 2001. Feeding habitsof the spotted hyaena in the Niokolo Koba National Park,
Senegal. Afr J Ecol 38:102–107.
6. Marchesi P, Marchesi N, Fruth B, Boesch C 2005. Census
and distribution of chimpanzees in Cote d’Ivoire. Primates
36:591–607.
7. Baldwin PJ, McGrew WC, Tutin CEG 1982. Wide-ranging
chimpanzees at Mt. Assirik, Senegal. Int J Primatol
3:367–385.
8. Plumptre AJ 2000. Monitoring mammal populations withline transect techniques in African forests. J Appl Ecol
37:356–368.
9. Bogart SL, Pruetz JD 2011. Insectivory of savanna chimpanzees (Pan troglodytes verus) at Fongoli, Senegal. Am J
Phys Anthropol 145:11–20.
10. Pruetz JD 2006. Feeding ecology of savanna chimpanzees(Pan troglodytes verus) at Fongoli, Senegal. In: Feeding
Ecology in Apes and Other Primates. Boesch C, Hohmann
G, Robbins MM (eds), Cambridge University Press,
Cambridge, pp. 161–182.

<NEWS>

Dr. Julius Keyyu Visits WRC
Gen'ichi Idani
Kyoto University, Japan(E-mail: gidani@wrc.kyoto-u.ac.jp)

Dr. Julius Key y u, Director of Research at the
Tanzania Wildlife Research Institute (TAWIRI), visited Japan in May 2012. He was invited by the Wildlife
Research Center (WRC), Kyoto University to attend aninternational symposium “Wildlife Studies in Tanzania”
and a workshop “Prospect and Cooperation for Wildlife
Research in Tanzania”.
The study of great apes by Japanese researchers in
Tanzania was pioneered by the late Dr. Kinji Imanishi andthe late Dr. Junichiro Itani in 1961. In 1965, the late Dr.
Toshisada Nishida was successful in provisioning wild
chimpanzees in the Mahale area, and Mahale Mountains
National Park was established in 1985 as the 11th National
Park in Tanzania.
Research in Mahale Mountains N. P. under the leadership of Dr. Nishida produced a lot of important results.
Key findings included the existence of the unit group(community), exchange of females among unit groups andthe patrilineal structure of chimpanzee society, fission andfusion of individuals within a unit group, male alliancesand dominant-subordinate relations, conflict between unitgroups, various sexual behaviors, infanticide and cannibalism, sharing behavior, cultural behaviors, including
tool-using behaviors. It can safely be said that these findings played an important role in developing our scientificunderstanding of a few differences between chimpanzees
and human beings.
At the same time as work was beginning at Mahale,
field research of wild chimpanzees was also carried out
in the Ugalla area, which is located 100 km north-east of
Mahale. Dr. Takayoshi Kano (now an honorary professorof Kyoto University) stayed alone in Ugalla during 19651967 and conducted a distribution survey of chimpanzees in Western Tanzania. In this survey, he established11
that the eastern limit for the distribution of chimpanzees
in Tanzania was at long. 31°1’ E on the left bank of the
Ugalla River, and the southern limit was at lat. 6°38’ S of
the Wansisi Hill. The left bank of the Ugalla River is also
the eastern limit of the distribution of the chimpanzee inAfrica. Unfortunately, research in Ugalla was not continued after that because efforts were concentrated on workat Mahale.
In 1994, however, Dr. Hideshi Ogawa (Chukyo
University) and I resumed the survey in Ugalla and studies there continue to the present day. Dr. Ogawa revised
the southern limit of the distribution of chimpanzees in
Tanzania to lat. 8°12’ S. We have also revealed uniqueecological characteristics of the miombo arid woodland of
Ugalla, and interesting studies of various mammals andraptors have been initiated there.
The program of the symposium held on 16th May,
2012 is shown below. It began with a keynote presentation by Dr. Keyyu, who introduced activities and researchin TAWIRI, and then the latest studies were described inthree presentations for each of Mahale and Ugalla. These
presentations dealt not only with chimpanzees, but also
referred to various other fauna, including raptors, and tothe vegetation and climate. The symposium stimulated
active discussion among many researchers and students.
We would like to express our sincere thanks to all of theparticipants in this symposium.

On the following day (the 17th May), a closed workshop was held among Dr. Keyyu and researchers of
Mahale and Ugalla. We discussed future collaboration
between TAWIRI and WRC in scientific research, conservation and education. Studies in most of the 16 nationalparks and 10 reserves for wild animals in Tanzania havebeen carried out by European and American researchers.
Japanese researchers have mainly promoted studies in
Mahale and Ugalla in western Tanzania. Although thereis the Katavi National Park in the south of Mahale, foreign researchers seldom work there. So we also discussed
a plan for Japanese researchers to develop studies in thewhole of western Tanzania, from Katavi N.P. to Mahaleand Ugalla. This workshop was a valuable event. One
excellent outcome was an agreement to form a MOU between TAWIRI and WRC in the near future.
Dr. Keyyu had friendly discussions with researchersand students at WRC, and he also visited Kyoto City Zoo
and Kyoto Aquarium. He was deeply touched to see aPan Africa News, 19(1), June 2012

12

<NEWS>

Book Launch

newborn gorilla in the zoo and wondered at the exhibits ofgreat salamanders and dolphins in the aquarium. Though
his stay in Kyoto was only 4 days, it was a fruitful time,and he left for home on 19th May. We hope that TAWIRIand WRC will develop more on each research and activity
in Tanzania.

<Symposium For Wildlife Studies In Tanzania>
Venue: Wildlife Research Center, Kyoto University
Schedule:
09:50 – Greeting, Prof. Gen'ichi Idani (WRC)10:00 – Keynote address, Dr. Julius Keyyu (TAWIRI),“Activities of TAWIRI and Research Topics”
11:15 – Dr. Michio Nakamura (WRC), “Long-Term
Research and Conservation of Chimpanzees at
Mahale”
12:15– Lunch time
13:20 – Dr. Noriko Itoh (WRC), “Long-Term Researchin Mahale: Climate, Vegetation, and Plant
Phenology”
14:10 – Dr. Koichiro Zamma (Great Ape ResearchInstitute, Hayashibara), “Populations of Mammals
and Chimpanzees in Mahale”
15:00 – Coffee break
15:10 – Ms. Midori Yoshikawa (WRC & Tokyo
University of Agriculture and Technology), “ThePresent States of Chimpanzees in Ugalla andOther Non-Protected Areas, Tanzania”
15:50 – Ms. Eriko Iida (WRC), “Wild Mammals in
Ugalla Area”
16:30 – Dr. Hiroshi Kaneda (WRC), “Ugalla, from
Eagle's View”
17:10 – Discussion
18:00 – Close
19:00 – Social gathering< Wo r k s h o p f o r P r o s p e c t a n d C o o p e r a t i o nfor W i l d l i fe Re s e a rch i n Ta n z a n i a (C l o s e d) >We discussed for future collaboration between
TAWIRI and WRC in scientific research, wildlife conservation and education.

Prof. Toshisada Nishida’s last book, Chimpanzees of
the Lakeshore: Natural History and Culture at Mahale(Cambridge University Press, 320 pp., 2012) got a launchparty at the annual meeting of the American Associationof Physical Anthropologists. The AAPA met in Portland,
Oregon, USA, from 11–14 April, 2012. The event was arranged by Martin Griffiths, commissioning editor for thebook, at Cambridge University Press. Conviviality was
guaranteed by CUP’s provisioning with a keg of craftedale from the hometown Rogue Brewery. Prof. William
McGrew proposed the toast to Prof. Nishida at the outset,
but later so many more persons arrived that another keg
had to be tapped and another toast was made. At least
200 persons, including many prominent primatologists,
attended, and the proceedings ended only with the drinking of the last drop of beer. It is hoped that Toshi (as heis affectionately known by his Western friends and colleagues) would have appreciated this celebration of hislife and achievements.

<BOOK INFO>

Long-Term Field Studies ofPrimates
Edited by Peter M. Kappeler and David P. Watts
Some primate field studies have been on-going for
decades, covering significant portions of individual life
cycles or even multiple generations. In this volume, leading field workers report on the history and infrastructureof their projects in Madagascar, Africa, Asia and SouthAmerica. More importantly, they provide summaries of
their long-term research efforts on primate behaviour,
ecology and life history, highlighting insights that were
only possible because of the long-term nature of the study.
The chapters of this volume collectively outline the many
scientific reasons for studying primate behaviour, ecology
and demography over multiple generations. This kind ofresearch is typically necessitated by the relatively slow
life histories of primates. Moreover, a complete understanding of social organization and behaviour, factors
often influenced by rare but important events, requires
long-term data collection. Finally, long-term field projects
are also becoming increasingly important foci of local
conservation activities.
Contents

Part I Introduction
1. The Values and Challenges of Long-Term Field Studies /Peter M. Kappeler, Carel P. van Schaik & David P. Watts

Part II Madagascar
2. Berenty Reserve, Madagascar: A Long Time in a SmallSpace / Alison Jolly

Pan Africa News, 19(1), June 201213

3. Beza Mahafaly Special Reserve: Long-Term Research on
Lemurs in Southwestern Madagascar / Robert W. Sussman,
Alison F. Richard, Joelisoa Ratsirarson, Michelle L.
Sauther, Diane K. Brockman, Lisa Gould, Richard Lawler& Frank P. Cuozzo
4. Long-Term Lemur Research at Centre Valbio, Ranomafana
National Park, Madagascar / Patricia C. Wright, Elizabeth
M. Erhart, Stacey Tecot, Andrea L. Baden, Summer J.
Arrigo-Nelson, James Herrera, Toni Lyn Morelli, Marina
B. Blanco, Anja Deppe, Sylvia Atsalis, Steig Johnson, Felix
Ratelolahy Chia Tan & Sarah Zohdy
5. A 15-Year Perspective on the Social Organization and Life
History of Sifaka in Kirindy Forest / Peter M. Kappeler &Claudia Fichtel

Part III America
6. The Northern Muriqui (Brachyteles hypoxanthus): Lessons
on Behavioral Plasticity and Population Dynamics from a
Critically Endangered Species / Karen B. Strier & Se´rgio L.
Mendes
7. The Lomas Barbudal Monkey Project: Two Decades of
Research on Cebus capucinus / Susan Perry, Irene Godoy& Wiebke Lammers

460 pages
Springer
Published January 2012
ISBN 978-3-642-22513-0(Hard Cover) 149,95 €ISBN 978-3-642-22514-7(eBook) 139,99 €8. Tracking Neotropical Monkeys in Santa Rosa: Lessons
from a Regenerating Costa Rican Dry Forest / Linda M.
Fedigan & Katharine M. Jack
9. The Group Life Cycle and Demog raphy of Brow n
Capuchin Monkeys (Cebus [apella] nigritus) in Iguazu´
National Park, Argentina / Charles Janson, Maria Celia
Baldovino & Mario Di Bitetti

Matsubara, Nobusuke Iwasaki & David S. Sprague

Part IV Asia
10. Social Organization and Male Residence Pattern in
Phayre’s Leaf Monkeys / Andreas Koenig & Carola Borries
11. White-Handed Gibbons of Khao Yai: Social Flexibility,
Complex Reproductive Strategies, and a Slow Life History/ Ulrich H. Reichard, Manoch Ganpanakngan & Claudia
Barelli

Part V Africa
12. The Amboseli Baboon Research Project: 40 Years ofContinuity and Change / Susan C. Alberts & Jeanne
Altmann
13. The 30-Year Blues: What We Know and Don’t Know
About Life History, Group Size, and Group Fission of Blue
Monkeys in the Kakamega Forest, Kenya / Marina Cords
14. Long-Term Research on Chimpanzee Behavioral Ecology
in Kibale National Park, Uganda / David P. Watts
15. Long-Term Field Studies of Chimpanzees at Mahale
Mountains National Park, Tanzania / Michio Nakamura &Toshisada Nishida
16. Long-Term Studies of the Chimpanzees of Gombe
National Park, Tanzania / Michael L. Wilson
17. Long-Term Research on Grauer’s Gorillas in Kahuzi-Biega
National Park, DRC: Life History, Foraging Strategies, andEcological Differentiation from Sympatric Chimpanzees/ Juichi Yamagiwa, Augustin Kanyunyi Basabose, John
Kahekwa, Dominique Bikaba, Chieko Ando, Miki

18. Long-Term Studies on Wild Bonobos at Wamba, Luo
Scientific Reserve, D. R. Congo: Towards the Understandingof Female Life History in a Male-Philopatric Species /Takeshi Furuichi, Gen’ichi Idani, Hiroshi Ihobe, Chie
Hashimoto, Yasuko Tashiro, Tetsuya Sakamaki, Mbangi N.
Mulavwa, Kumugo Yangozene & Suehisa Kuroda

Part VI Summary
19. Long-Ter m, Individual-Based Field St udies / Tim
Clutton-Brock

Index<BOOK INFO>The Ecological Impact of LongTerm Changes in Africa's Rift
Valley (Environmental Science,
Engineering and Technology)
Edited by Andrew J. Plumptre
Despite Africa’s rich biodiversity and the importanceof its ecosystem services, it has relatively few collaborative, network-based studies that examine the ecologicalimpacts of climate change. This book marks the begin-Pan Africa News, 19(1), June 201214
ning of such a collaboration. It covers ecological information that spans across five countries in the Albertine Riftregion, reflects over 50 years of research, and includes
contributions from 65 researchers who represent 44 organizations at work in 11 sites. It provides invaluable
information about past and current trends in the status of
species, ecosystems and associated threats, as well as recommendations for interventions.

6. The Effects of Environmental and Anthropogenic Changes
on the Savannas of the Queen Elizabeth and Virunga
National Parks / Andrew J. Plumptre, Derek Pomeroy,
Jared Stabach, Nadine Laporte, Margaret Driciru, Grace
Nangendo, Frederick Wanyama & Aggrey Rwetsiba
7. Long-Term Ecological and Socio-Economic Changes in andaround Bwindi Impenetrable National Park, South-Western
Uganda / Aventino Kasangaki, Robert Bitariho, Phil Shaw,
Martha Robbins & Alastair McNeilage
8. Long Term Changes in the Virunga Volcanoes / Katie
Fawcett, Glenn Bush, Anton Seimon, Guy Picton Phillips,
Deo Tuyisingize & Prosper Uwingeli
9. Long Term Changes in a Montane Forest in an Region ofHigh Human Population Density / Nerissa Chao, Felix
Mulindahabi, Julian Easton, Andrew J. Plumptre, Anton
Seimon, Adrian Martin & Robert Fimbel
10. Long-Term Changes in Habitats and Ecology of African
Apes in Kahuzi-Biega National Park, Democratic Republicof Congo / Juichi Yamagiwa, Augustin K. Basabose, John
Kahekwa, Dominique Bikaba, Miki Matsubara, ChiekoAndo, Nobusuke Iwasaki & David S. Sprague
11. Long-Term Changes in the Ecological Factors Surrounding
the Chimpanzees of the Gombe National Park / Lilian
Pintea, Anne Pusey, Mike Wilson, Ian Gilby, Anthony
Collins, Shadrack Kamenya & Jane Goodall

308 pages
Nova Science Pub Inc
Published March 30, 2012
ISBN-10: 1611227801
ISBN-13: 978-1611227802
Hardcover: $145.00

12. L o ng-Te r m C h a nge s i n t h e So cia l a n d Na t u r a l
Environments Surrounding the Chimpanzees of the Mahale
Mountains National Park / Noriko Itoh, Michio Nakamura,
Hiroshi Ihobe, Shigeo Uehara, Koichiro Zamma, Lilian
Pintea, Anton Seimon & Toshisada Nishida
13. Long Term Changes in Africa’s Western Rift Valley:
Synthesis of Main Findings / Andrew J. Plumptre

Index

Contents

Contributors
Foreward / Elizabeth Chadri
Preface / Andrew J. Plumptre
1. Africa’s Western Rift: An Introduction / Andrew J.
Plumptre & Cecily Kabagumya
2. Regional Climatology of the Albertine Rift / Anton Seimon& Guy Picton Phillipps
3. Environmental and Anthropogenic Changes in and around
Budongo Forest Reserve. / Fred Babweteera, Douglas Sheil,
Vernon Reynolds, Andrew J. Plumptre, Klaus Zuberbuhler,
Cathrine M. Hill, Amanda Webber & Mnason Tweheyo
4. Long Term Changes at Toro-Semliki Wildlife Reserve /Randy Patrick, Deborah Patrick & Kevin D. Hunt
5. Complex Responses to Climate and Anthropogenic
Changes: An Evaluation Based on Long-Term Data from
Kibale National Park, Uganda / Colin A. Chapman, Lauren J.
Chapman, Ria Ghai, Joel Hartter, Aerin L. Jacob, Jeremiah
S. Lwanga, Patrick Omeja, Jessica M. Rothman & Dennis
Twinomugisha

Pan Africa News, Vol. 19, No.1
Published in June, 2012
Address: c/o Human Evolution Studies,
Dept. of Zoology, Faculty of Science,
Kyoto Univ., Kyoto, 606-8502, JAPANTEL: (+81)75-753-4093
FAX: (+81)75-753-4115
E-mail: pan.editor@gmail.com
URL: http://mahale.main.jp/PAN/
ISSN: 1884-751X (Print), 1884-7528 (Online)