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

DECEMBER 2008

VOL. 15, NO. 2

P. A. N. EDITORIAL STAFF

Contents

Chief Editor:
Toshisada Nishida, Japan Monkey Centre, Japan<NOTE>
Fongoli chimpanzee (Pan troglodytes verus) eats banded
mongoose (Mungos mungo)
S.L. Bogart, J.D. Pruetz & D. Kante
15

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<NOTE>
Population density of chimpanzees in Tanzania
M. Yoshikawa, H. Ogawa, T. Sakamaki & G. Idani

17<NOTE>
Arbors and cuttings: New trials for Green Corridor Projectat Bossou-Nimba
G. Ohashi, R. Hasegawa, M. Kourouma& T. Matsuzawa
20

Editorial Secretaries:
Agumi Inaba, Japan Monkey Centre, Japan
Noriko Itoh, Japan Monkey Centre, Japan
Koichiro Zamma, GARI Hayashibara, Japan

Instructions to Authors:
Pan Africa News publishes articles, notes, reviews, forums,
news, essays, book reviews, letters to editor, and classified ads(restricted to non-profit organizations) on any aspect ofconservation and research regarding chimpanzees (Pantroglodytes) and bilias (Pan paniscus). Contributors arerequested to write in English and the papers except forums,
reviews and essays should usually be 1,500 words or less.
Articles and notes will be peer-reviewed by at least one
appropriate expert on request of the PAN editorial staff.
Manuscripts should be formatted as DOC or RTF files andsubmitted by e-mail to: pan.editor@gmail.com
Photos and figures should be formatted as JPEG or GIF files
and sent separately by e-mail attachments.

<NOTE>
Newly observed predation of wild birds by M-groupchimpanzees (Pan troglodytes schweinfurthii) at Mahale,
Tanzania
M. Fujimoto & M. Shimada
23<NOTE>The Ebo Forest: Four years of preliminary research andconservation of the Nigeria-Cameroon chimpanzee (Pantroglodytes vellerosus)
E.E. Abwe & B.J. Morgan
26<NOTE>
Coprophagy by the semi-habituated chimpanzees of Semliki,
UgandaC.L.R. Payne, T.H. Webster & K.D. Hunt
29

Publication will be made twice a year in June and December.
Deadline for manuscripts is one month before publication (i.e.
the ends of May and November).

<NOTE>

1. Department of Anthropology, Iowa State University

Fongoli chimpanzee (Pantroglodytes verus) eats banded
mongoose (Mungos mungo)
1,2

1,2

S.L. Bogart , J.D. Pruetz

3and D. Kante

2. Ecology and Evolutionary Biology Interdepartmental
Program, Iowa State University
3. Fongoli Savanna Chimpanzee Project, Kedougou, Senegal

INTRODUCTION
We provide the first evidence of a chimpanzee(Pan troglodytes verus) preying upon a banded15

16
mongoose (Mungos mungo). A subadult female wasobserved consuming a banded mongoose at the Fongolisavanna chimpanzee study site, Senegal. This recording
provides new data and adds a small carnivore to the listof animal prey known for the chimpanzee diet.
Banded mongooses are found throughout
sub-Saharan, central, and eastern Africa1. They areusually found in savanna woodland, habitually staying
near water sources1, but they are found in open savanna
habitats at Fongoli, Senegal. The banded mongoose
resides in groups greater than 35 individuals, which retire
at night into dens, usually located in termite mounds1.
The banded mongoose diet consists mainly of insects,
worms, larvae, and small vertebrates1.
The Fongoli study site is located in southeastern
Senegal (12o39’ N, 12o13’ W), and the chimpanzee
community has been observed since 2001. The numberof chimpanzees fluctuates annually, but 33–36
individuals usually comprise the community. The
chimpanzees at Fongoli are semi-habituated to humanobservers in that some of the adult females are still
nervous in the presence of observers. The community hasa home range of more than 63 square kilometers2.
Fongoli is a mosaic environment including both open
and closed canopy habitats, such as woodland andgallery forest2. Chimpanzees here are similar to thosestudied elsewhere in that their diet can be consideredomnivorous, containing everything from seeds to otheranimals3. In general, the chimpanzee diet is comprised
mostly of fruit (60%) with animal prey accounting for
8% (averaged from 12 sites including Fongoli)2. Animal
prey does not constitute a high proportion of thechimpanzee diet. However, it can provide essential
nutritional components that vegetation resources lack3,4.
OBSERVATIONThe banded mongoose at Fongoli are observed on
average in every month of the year and range in groupsize from two to 24 individuals with a mean group sizeof 5.6 individuals (N=203 individuals, 36 sightings over
22 months from 2005–2008) and found in open savanna
and closed forested habitats within the year of this study.
On August 20, 2006, a group of more than five banded
mongooses were seen on the plateau at Djendji waterhole at 1503 hours; they then traveled into the gallery

Pan Africa News, 15(2), December, 2008
forest. Several days later, August 25, 2006, at the same
locale, a chimpanzee was observed feeding on amongoose.
At 1620 hours, Tia, a sub-adult female estimated to
be about 13 years old, obtained an animal that did notresemble any of the three monkey species known at
Fongoli. The capture was not witnessed so it is notknown how she acquired the animal. Enough of theprey’s body was intact to determine that the kill wasrecent and to allow identification of the animal as a
banded mongoose. At one point, she did give somepieces of meat to a sub-adult male. Tia fed on themongoose until 1636 hours, at which time she moved outof sight. When we located Tia again at 1650 hours(12°40.069N, 12°11.130W), the prey’s head was gone,
but the black dorsal stripes that characterize the banded
mongoose were still visible. Tia ate the meat with leaves
from an unidentified tree until 1750 hours when she
again moved out of sight. When we attempted to follow
her, Tia could not be found again within the group. The
group made beds at 1820 hours; we attempted to find Tia
the next morning in an effort to find the carcass of themongoose, but we were unsuccessful.
There has never been any record of mongoose in
the diet of the Fongoli chimpanzee, in feces nor in
observation. Additionally, there has never been any
report of the banded mongoose as a prey item at anyother chimpanzee site in Africa, where banded
mongooses are known to exist (Assirik, Senegal; Gombeand Mahale, Tanzania; Kibale, Uganda)5. Elsewhere in
Tanzania, chimpanzees have been recorded to consume
the white-tailed mongoose (Ichneumia albicauda)6.
DISCUSSION
This instance of mongoose eating is anecdotal7,8,
but continuous study may reveal that this prey item ismore common. Given that adult males are targeted forstudy, and certain females are relatively shy aroundhuman observers, further evidence is needed todetermine whether the banded mongoose is a regularfood source for the Fongoli chimpanzees or if it is an
opportunistic prey species. Most likely this occurrence
was the result of an opportunistic hunting event by the
sub-adult female, since it has never been observed until
now. This community has been observed hunting thePan Africa News, 15(2), December, 2008
nocturnal lesser bushbaby (Galago senegalensis) withfashioned tools9, and Tia ranks among the top five
bushbaby hunters in the community. Since the capture of
the mongoose was not observed it cannot be said whether
a tool was used or not. However, given the behavior of
the mongoose relative to a Galago1, there is no reason to
presume that tools were used. The capture of themongoose may have been similar to techniques used inacquiring monkeys or even squirrels, which are chased
along branches or the ground (Pruetz, unpublished data).
This is the first report of a chimpanzee feeding on abanded mongoose. Our findings may be an example ofcross-cultural differences in diet among chimpanzee
communities3,10, but this hypothesis has yet to be tested
with reference to the Fongoli chimpanzees. The inclusionof the banded mongoose in the diet may also be due toenvironmental constraints, such as the lack of mongooses
at some of the other chimpanzee sites. Moreover, thefaunal prey sources at Fongoli are limited to only threemonkey species (Papio hamadryas papio, Chlorocebus
aethiops, Erythrocebus patas), one nocturnal prosimian,
bushbucks(Tragelaphus
scriptus),squirrels(Heliosciurus gambianus), and warthogs (Phacochoerus
aethiopicus). The Fongoli chimpanzees have been
observed to pursue all but the warthog. The absence of
red colobus (Procolobus badius), one of the main preyspecies at many chimpanzee sites, may explain the low
occurrence of meat eating at Fongoli (5.4% of diet)2,9. It
also may be that the low variety of prey species leads tothe consumption of other small mammals, like thebanded mongoose, at Fongoli.
Acknowledgements. S. Bogart thanks Wenner-Gren Foundation,
Individual Research Grant for funds. J. Pruetz was supportedby The Leakey Foundation and the National Geographic
Society. We thank M. Camera, A. Quinn, J. Quinn, and S.
Macdonald for assistance in the field and A. Quinn forcomments on the manuscript.

17
Cambridge, pp. 159–179.
3

McGrew WC 1983. Animal foods in the diets of wild
chimpanzees (Pan troglodytes): why cross-culturalvariation? J Ethol 1: 46–61.

4

Sussman RW 1987. Species-specific dietary patterns inprimates and human dietary adaptations. In: The Evolutionof Human Behavior: Primate Models, Kinzey WG (ed),State University of New York Press, Albany, pp. 143–156.

5

Russak SM, McGrew WC 2008. Chimpanzees as fauna:comparisons of sympatric large mammals across long term
study sites. Am J Primatol 70: 402–409.

6

Goodall J 1986. The Chimpanzees of Gombe: Patterns of
Behavior. Belknap Press, Cambridge MA.

7

Sarringhaus LA, McGrew WC, Marchant LF 2005. Misuseof anecdotes in primatology: lessons from citation analysis.
Am J Primatol 65: 283–288.

8

Bates LA, Byrne RW 2007. Creative or created: using
anecdotes to investigate animal cognition. Methods 42:
12–21.

9

Pruetz JD, Bertolani P 2007. Savanna chimpanzees, Pantroglodytes verus, hunt with tools. Curr Biol 17: 412–417.

10 Boesch C, Gone Bi ZB, Anderson D, Stahl D 2006. Foodchoice in Taï chimpanzees: are cultural differences present?
In: Feeding Ecology in Apes and Other Primates, Hohmann
G, Robbins MM, Boesch C (eds), Cambridge University
Press, New York, pp. 183–201.

<NOTE>

Population density ofchimpanzees in Tanzania
Midori Yoshikawa1, Hideshi Ogawa2,
Tetsuya Sakamaki3 and Gen’ichi Idani4
1. Great Ape Research Institute (GARI), Hayashibara
2. School of International Liberal Studies, Chukyo University
3. Faculty of International Studies, Meiji-Gakuin University
4. Wildlife Research Center, Kyoto University

REFERENCES
1

2

Estes RD 1991. The Behavior Guide to African Mammals:
Including Hoofed Mammals, Carnivores, Primates.
University of California Press, Berkeley.
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,

INTRODUCTION
Chimpanzees (Pan troglodytes) are distributed
around the eastern shore of Lake Tanganyika in western
Tanzania. Based on an extensive survey conducted in the
1960s, Kano1 showed that chimpanzees were distributed

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in (a) Gombe, (b) Lilanshimba, (c) Ugalla, (d) Masito,(e) Mukuyu, (f) Mahale, (g) Karobwa, and (h) Wansisi(Fig. 1). In 1996, Ogawa et al.2 identified anotherchimpanzee habitat, (i) Lwazi (Fig. 1). However,chimpanzee population density and distribution in
Tanzania have not been revised since Kano’s report1. In
this paper, we showed the location in which we foundbeds and other traces of chimpanzees, and estimatedchimpanzee population densities in (d) Masito, (e)Mukuyu, and (i) Lwazi. Besides, with the use of some
additional unpublished data, we revised estimatedchimpanzee population densities in (b) Lilanshimba3, (c)Ugalla4, (g) Karobwa5, and (h) Wansisi6 using the
DISTANCE 5.0 computer program7.
METHODSIn Tanzania, most chimpanzee habitats are savanna
woodlands (miombo woodlands) dominated bydeciduous trees of Brachystegia and Julbernardia, with asmall proportion of evergreen riverine forests, grasslands,
and bamboo thickets.
We visited all the areas indicated in Fig. 1 (a–i)
and other areas where chimpanzees might have been
distributed, using 1/50000 topographical maps,
vegetation maps, satellite data, and information
provided by locals. In addition to the areas reported
in previous studies2,3,4,5,6 and Table 1, Ogawa andKanamori visited the Ufipa Escarpment (8°15’S,
32°00’E to 8°30S, 32°15’E) from Aug. 9–13 andAug. 31–Sept. 8, 2001. Ogawa visited (a) Gombe
from Sep. 3–6, 2003 and Mishamo from Aug.4–9,
2006. Sakamaki made an extensive survey in (f)Mahale from Jan. 23–30, 2006. Idani and Sakamaki
stayed at the area between (g) Karobwa and (h)Wansisi from Jan. 23 to 30, 2006. Ogawa stayed at
Mishamo (5°36’S, 30°28’E) from Aug. 4 to 9, 2006.
We conducted line transect bed censuses at (d)Masito: Lemera (5°15’S, 30°20’E) and Kasakatisites (5°20’S, 29°55’E), (e) Mukuyu: Lungonya site
(5°40’S, 29°55’E), and (i) Lwazi: Msalamba,
Mwinbi, Luanji, and Senge sites (8°10–25’S,
31°00–15’E)2. With the use of some additional
unpublished data, we revised estimated chimpanzee
population densities at (b) Lilanshimba3, (c)Ugalla4: Bhukarai, Bukombe, Issa, Issa Basin,

Pan Africa News, 15(2), December, 2008
Mfubasi, Mfuwazi, Mtongesi, Nguye, Nkondwe, andShangwasites; (g) Karobwa5: Ntakata and Kapalagulu5sites; and (h) Wansisi6.
We measured the distance of some census lines
using a tape measure; for other reconnaissance path, weused a portable GPS (Garmin 60CSx) and/or a pedometer.
We walked not only on traditional paths used by locals,
but also on paths used by wild animals, as well as off
paths, and traveled through various types of vegetation.
We measured the perpendicular distance from the census
line to a chimpanzee bed using a distance meter (Nikon
Laser 800S), counting the steps of local guides, or by eye
after being trained to estimate distance in this way.
Densities of chimpanzee beds were estimated using
the DISTANCE 5.0 computer program7. We used the
Akaike Information Criteria (AIC) selection rule tochoose a model from among three key detection
functions (uniform, half-normal, hazard-rate) and threeadjust detection functions (cosine, simple-polynomial,and Hermite-polynomial). We used those bed densities toestimate densities of bed-building chimpanzees,

Fig. 1. Distribution of chimpanzees in Tanzania.

Pan Africa News, 15(2), December, 2008

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Table 1. Population density of chimpanzees outside of national parks in Tanzania.

assuming that chimpanzees build 1.15±0.0478 bed perday. We used 260 days4 as the mean lifespan of achimpanzee bed, because previous research has shownthat bed trees in (c) Ugalla last for more than 260 days,
and the study areas had climate, vegetation, and bed trees
similar to Ugalla, with the exception of Gombe (a) and
Mahale (f). We also conducted a marked bed countcensus at the Nguye site in Ugalla in 2008.
RESULTS
Fig. 1 shows the distribution of chimpanzees inwestern Tanzania. We confirmed that chimpanzees
continue to inhabit all the areas they had inhabited in the
1960s. In addition to these areas, we found chimpanzee
beds between (h) Wansisi and (g) Karobwa (6°17’S,30°22’E). In contrast, we found no evidence ofchimpanzees in flat areas, such as the Mishamo Basin.
As reported by Kano1, chimpanzees mainly seem toinhabit mountainous areas.
Table 1 summarizes chimpanzee population density
outside national parks in Tanzania. In all areas, densities
were lower in the 1990s–2000s than in the 1960s1, withthe exception of (c) Ugalla (Wilcoxon signed rank test, n= 6, z = -1.99, p < 0.05).
Our marked bed count census at the Nguye site in
Ugalla revealed that 36 beds were made during 180 days
along five 5-km transect lines, indicating a density of
0.09 individuals/km2 (95% Confidence interval

0.05–0.24). This estimation is not contradictory to theassumption that chimpanzee beds can last as long as 260
days in this area, because the results of the standing crop
bed count census in Ugalla also revealed a density of
0.10 individuals/km2, when we used 260 days as themean lifespan of a chimpanzee bed (Table 1).
DISCUSSION
Kano1 hypothesized that the chimpanzee populationin (h) Wansisi was isolated from chimpanzees in (g)Karobwa. However, we observed chimpanzee beds
between these two areas. The hilly geographical features
and type of vegetation suggest that the chimpanzee
habitat of the Wansisi area may be narrowly connectedwith that of the Karobwa area. If so, the areas shown in(c–h) may form one continuous huge chimpanzee habitat.
No researchers have observed bed-building activity in (i)Lwazi, but DNA sequences from fecal samples haveindicated that the inhabitants are chimpanzees (Pantroglodytes) and may either be or be close to Pan
troglodytes schweinfurthii (Tashiro, unpublished data).
To date, our surveys have confirmed as many as four
local chimpanzee populations in Tanzania, in: (1) Gombe,(2) Lilanshimba, (3) Ugalla-Masito-Mukuyu-MahaleKarobwa-Wansisi, and (4) Lwazi (these populations do
not include the introduced population on Rubondo Islandin Lake Victoria). Our surveys indicate that no local
populations have been exterminated during the past 50

20
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years.
However, it is likely that each chimpanzee habitat
area has shrunk. Calculation of the possible and actual
chimpanzee habitat areas will require additional surveys
and analyses using satellite and GIS data. The number ofchimpanzees has clearly been reduced because
population densities in each area were lower than in the
1960s1. Although direct comparison is difficult due to thediffering methods of estimation, Table 1 indicates thatthe population density has been reduced since the 1960s.
Density has been markedly reduced in (b) Lilanshimba,(e) Mukuyu, (g) Karobwa, and (h) Wansisi, probably due
to the large human population in these areas, whereas
density was unchanged in (c) Ugalla. In (b) Lilanshimba,(e) Mukuyu, (g) Karobwa, (h) Wansisi, and (i) Lwazi,
increasing numbers of immigrant farmers have cultivatedfields, except in steep mountainous areas. Trees havebeen cut down for use as lumber and firewood.
Burundian refugees at the Mishamo settlement set manysnares in (e) Mukuyu and (g) Karobwa, and Congolese
refugees set snares in (b) Lilanshimba; this poaching
resulted in deaths of chimpanzees in these areas3,5,6.
Proper conservational planning is vital to protect
chimpanzees and their habitats in Tanzania.
Acknowledgements． We are grateful to T. Kano, M. Kanamori,
H. Y. Kayumbo, E. T. Massawe, J. Itani, J. Moore, and T.
Nemoto for their help in the field work. This study waspermitted by the Tanzanian authorities (COSTECH and
TAWIRI) and was financially supported by a Grand-in-Aid for
Scientific Research from MEXT, Japan (09041160; 06061064;
1257597; 17255005), Global Environment Research Fund
(F-061 to T. Nishida) of the Ministry of the Environment,
Japan.

REFERENCES
1

Kano T 1972. Distribution and adaptation of thechimpanzee on the eastern shore of Lake Tanganyika.
Kyoto Univ Afr Stud 7: 37–129.

2

Ogawa H, Kano T, Kanamori M, Massawe, E. 1999.
Chimpanzee habitat in the southwestern Rukwa region,
Tanzania. Primate Res 15: 147–151.

3

Ogawa H, Sakamaki T, Idani G 2006. The influence ofCongolese refugees on chimpanzees in the Lilanshimba
area, Tanzania. Pan Afr News 13: 21–22.

4

Ogawa H, Idani G, Moore J, Pintea L, Hernandez-Aguilar
A. 2007. Sleeping parties and nest distribution ofchimpanzees in the savanna woodland area, Ugalla,
Tanzania. Int J Primatol 28: 1397–1412.

Pan Africa News, 15(2), December, 2008
5

Ogawa H, Moore J, Kamenya S 2006. Chimpanzees in the
Ntakata and Kakungu areas, Tanzania. Primate Conserv 21:
97–101.

6

Ogawa H, Moore J, Kanamori M, Kamenya S 2004. Reporton the chimpanzees of the Wansisi and Makomayo areas,
Tanzania. Pan Afr News 11: 3–5.

7

Buckland ST, Anderson DR, Burnham KP, Laake JL 1993.
Distance Sampling: Estimating Abundance of Biological
Populations. Chapman and Hall, London.

8

Plumptre AJ 2000. Monitoring mammal populations withline transect techniques in African forests. J Appl Ecol 37:
356–368.

<NOTE>

Arbors and cuttings: New
trials for Green Corridor
Project at Bossou-Nimba
Gaku Ohashi1,Ryo Hasegawa2,
Makan Kourouma3 and Tetsuro Matsuzawa1
1. Primate Research Institute, Kyoto University
2. Phytoculture Control Co., LTD.
3. Institut de Recherche Environnementale de Bossou

Since 1976, wild chimpanzees have been studied at
Bossou, Guinea, in West Africa1,2. The Bossou groupitself is extremely endangered. Since the beginning of
our study more than 30 years ago, no female chimpanzee
immigration has been recorded, whereas all of the female
chimpanzees born at Bossou disappeared around sexual
maturation3. As a result, the percentage of aged
individuals is increasing in the group. To make mattersworse, the number of Bossou chimpanzees suddenly
decreased to 12 in 2003, due to an epidemic ofrespiratory disease4. For the group’s survival, individual
immigration from the nearest groups is essential.
About 10 km away from Bossou, at least one
chimpanzee group lives in the Nimba Mountains5. Inorder to promote individual interchange between the
Bossou and Nimba groups, we started in 1997 the“Green Corridor Project”, which has involved planting
trees along a 4 km long expanse of savanna area
separating Bossou from Nimba6. Owing to this effort, we
can see today small forest patches growing in this area7.

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Pan Africa News, 15(2), December, 2008
During a decade, we have grown saplings in a tree
nursery at Bossou, and transported them into the savanna.
During transplantation, the environment around thesaplings is drastically changed. The saplings are forced
to receive strong sunshine in the savanna. For this reason,
the saplings often died within a few days. To minimize
damage caused by solar insolation, we started to makearbors in the savanna to protect young saplings. The
arbors were constructed from natural materials, as local
people do for their temporal ceremonies: we usedbamboos as columns, leaf stalks of Raffia as beams, andpalm leaves as roof. The roof allows the passage of water,whilst providing shade.
On September 2007, we constructed 3 arbors andtransplanted 25 Uapaca heudelotti saplings under each ofthem (i.e., 75 saplings in total). One year later, wechecked the condition of the Uapaca trees under thearbors. Whereas 13 trees were damaged by termites, theother 62 trees survived and thrived (82.7%). Some trees
grew to 3 cm in diameter and 130 cm in height (Fig. 1).
The arbor itself was small, but it proved to be a usefulapproach for creating patches of forest in the center ofthe savanna, compared to direct transplantation without
protection which was often challenging. We continued toconstruct arbors in 2008 after the first trial, and a total of
23 arbors are now protecting young trees against strong
sunshine of the savanna.
In 2007, we tested another planting method using
tree cuttings. When we checked fences around villages,
we noticed that some sticks were sprouting. We
identified the sticks used for fences in 3 villages,
including Bossou, and checked which species had shoots.
Our study led us to investigate 8,998 sticks; 176 sticks
were found to have new buds or leaves (Table 1). We
counted 579 sticks of Uapaca heudelotti, but they did nothave any shoots. When we transported saplings from tree
nursery into the savanna, Uapaca was one of the mostsuccessful ones7. However, the Uapaca may be
unbefitting in the tree cutting method. On the other hand,
fifty one out of the 176 sprouting sticks were Spondias
cytherea. We collected 1,523 cuttings of Spondias
cytherea from forest, and directly planted the cuttings
around gallery forest and small forest patches in savanna.
Only 3 weeks later, 891 out of the 1,523 (58.5%) cuttings
were found to have produced new shoots (Fig. 2). We

Fig. 1. Uapaca heudelotti trees under thearbor. They grew up to 130 cm in height.

Fig. 2. Cutting of Spondias cytherea planted
around patchy forest.

still need to monitor growth development of these tree
cuttings over time, but this method may be useful forenlarging the forest patches adjoining the savanna.
These new attempts sparked great interest in thelocal people, because they consist of using techniques
already familiar to them and applied to a conservation
oriented reforestation project. Some villagers repaired
the arbors voluntarily, while others started to plantcuttings including other species voluntarily. Changing
savanna to forest by using the local methods is expected
to increase environmental awareness of local
communities and increasing their participation inconservation efforts.

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Pan Africa News, 15(2), December, 2008
Table 1. Germination rate of cuttings. We identified the sticks used for fencing inthree villages, including Bossou, and checked which species had produced shoots.

Acknowledgements. This work was financially supported by
MEXT-07102010, 12002009, and 16002001 and JSPS-HOPE
to TM. The current project is supported by the Global
Environment Research Fund (F-061) of the Ministry of theEnvironment, Japan to Dr. Toshisada Nishida. We thank theresearch assistants for promoting this project such as Boniface
Zogbira, Paquille Cherif, Pascal Goumy, Jules Dore, and Henry

Gberegbe. Thanks are also due to the Guinean authorities, Drs.
Kabine Kante and Tamba Tagbino from the DNRST. We arealso grateful for the effort of colleagues who contributed thisproject during various stages; Drs. Yukimaru Sugiyama, Gen
Yamakoshi, Tatyana Humle, and Nicolas Granier.

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Pan Africa News, 15(2), December, 2008
REFERENCES
1

Matsuzawa T 2006. Bossou 30 years. Pan Afr News 13:
16–19.

2

Sugiyama Y, Koman J 1979. Social structure and dynamics
of wild chimpanzees at Bossou, Guinea. Primates 20:
323–339.

3

Sugiyama Y 2004. Demographic parameters and life
history of chimpanzees at Bossou, Guinea. Am J Phys
Anthropol 124: 154–165.

4

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

5

Koops K, Matsuzawa T 2006. Hand clapping by achimpanzee in the Nimba Mountains, Guinea, West Africa.
Pan Afr News 13: 19–21.

6Hirata S, Morimura N, Matsuzawa T 1998. Green passage
plan (tree-planting project) and environmental education
using documentary videos at Bossou: A progress report.
Pan Afr News 5: 18–20.

7Matsuzawa T 2007. Assessment of the planted trees inGreen Corridor Project. Pan Afr News 14: 27–29.

<NOTE>

Newly observed predation ofwild birds by M-groupchimpanzees (Pan troglodytes
schweinfurthii) at Mahale,
Tanzania
Mariko Fujimoto1 and Masaki Shimada1,2
1. Graduate School of Human Cultures, The University of Shiga
Prefecture
2. Japan Society for the Promotion of Science

INTRODUCTION
A number of predatory habits of wild chimpanzees(Pan troglodytes) have been reported across Africa1,2,3,4.
Most of these studies have reported mainly meat-eatingof mammals. There are a few descriptions of predation of
birds by chimpanzees both direct and indirect evidence
in some study sites4-11.
When chimpanzees hunt and eat primates of otherspecies, most chimpanzees present actively join in thehunt and the subsequent meat-eating1,2,12. In contrast,
when preying on non-primate mammals, such as blue
duiker, bush pig, and squirrel and so on, chimpanzees
hunt in an opportunistic manner and not in groups3,12,13.
Few studies have described predation on wild birds bychimpanzees. This study reports 3 direct observations ofmeat-eating of birds by chimpanzees in Mahale.

Study site and observation method
The M-group chimpanzees in Mahale have been
observed for more than 40 years and all members havebeen identified and well habituated to human observers.
The first author, MF, conducted the field studies between
May 2005 and April 2006. The second author, MS,
conducted his investigation between September 2001 and
September 2002. We observed the chimpanzees in detailwhen they engaged in predatory behavior.
RESULTS
Case 1On October 8, 2001, at 1233, MS observed 2 adultfemales TZ (age, 19 years) and NK (age, 31 years) up ina tree. TZ put her hand in the hole of the trunk andretrieved a nestling of a trumpeter hornbill (Ceratogymna
bucinator). NK then approached TZ and took thenestling from TZ and moved to a spot approximately 3
meters away. At 1235, an adult female JN (estimated age,
27 years) inserted her hand into the same hole but did not
find anything. NK held on to the carcass but did not eat
it; she left the carcass on the ground at 1237. Thereafter,
TZ climbed down to the ground and picked the carcass
up. At 1240, she started eating the head of the carcass.
An adolescent female ZL (estimated age, 15 years)
approached TZ and uttered a pant-grunt. TZ stamped herfoot while holding the carcass in her mouth, and ZL
screamed and fled. At 1249, NK and TZ went away. MS
did not witness whether TZ abandoned the carcass before
moving or not.
Case 2On August 2, 2005, at 1434, MF observed TZ (age,23 years) climb a tree and walk slowly on one of its
branches. MF did not clearly witness the moment when
TZ procured something from a bird’s nest, but observed

24
16
that at 1438, 2 parent white-necked ravens (Corvus
albicollis) attacked TZ. TZ then descended to a lower
spot on the tree, and started eating the nestling at 1508.
At 1525, TZ’s son TD (age, 4 years) stared at TZ’s mouth
and approximated his mouth to his mother’s andobtained a small piece of meat. An adult female AK(estimated age, 24 years) also approached and stared at
TZ for a few minutes, and then reached for the carcass inTZ’s hand. AK succeeded in obtaining a piece of meat at
1548 and ate it with her daughter AC (age, 7 years). At
1551, an adult female SY (estimated age, 22 years) and
her daughter SW (age 2 years) appeared. SY groomed
TZ for about 3 minutes. Thereafter, SY repeatedly looked
into TZ’s mouth and hand, and then, TZ groomed SY for
3 minutes. TZ restarted eating the carcass, and SY
extended her hand toward TZ’s hand. SY acquired asmall piece of meat at 1601. TZ and TD started moving
and abandoned the carcass at 1609.

Case 3On November 1, 2005, MF observed chimpanzees
preying on a red colobus and consuming its meat for
about one hour. At 1322, an adult male DW (age, 17
years) was observed holding a carcass of an adult black
kite (Milvus migrans). At 1325, MF’s research assistantnoted that an adolescent male PR (age, 14 years) also
held a carcass of a black kite. Both DW and PR departed
and were quite a distance away from the other members.
When the group engaged in eating the red colobus began
to disperse at 1540, a juvenile male OS (age, 7 years)
arrived holding the body of an adult black kite. He
carried and started eating it by himself. His mother OP(estimated age, 34 years), his sister RB (age, 19 years),and RB’s daughter RC (age, 7 years) approached OS, andhe allowed them to eat the kite. Afterward, two adultfemales got small pieces of meat from OS. The alphamale AL (age, 17 years) then arrived and stared at OS but
left quickly. After this, we lost OS momentarily, andsubsequently saw his niece RC with the carcass (Fig. 1).
Then, OS took and carried it. At 1713, OS abandoned thecarcass and started moving with the other groupmembers. We were unable to determine whether OS had
simply picked up one of the carcasses that PR or DW hadabandoned or whether he captured it independently. We
can report that at least 2 black kites were hunted andeaten by chimpanzees.

Pan Africa News, 15(2), December, 2008

Fig. 1. A juvenile female with a carcass of a blackkite.

DISCUSSION
Our study revealed some characteristics of bird
predation by chimpanzees based on 3 direct observations
and some information about the predation of wild birds
by chimpanzees obtained through previous studies (Table
1).
First, capture of nestlings from their nests seems
similar to that of non-primate mammals in that it issolitary and opportunistic. Our observations showed thatchimpanzees do not hunt birds in groups. In the 1st and2nd episodes, TZ took a nestling from a bird’s nest. She
approached each nest and took the nestling, which
offered no resistance to the predator. The bird parents
attacked TZ in the 2nd case, but she succeeded in
capturing the nestling.
Second, we can point out individual differencesamong chimpanzees even within a unit group in theirtendency to hunt and eat birds. Not all the chimpanzees
present in the 3 episodes begged for meat actively, while
the hunter and meat possessor ate the meat. In the 1stcase, although the adult female NK appropriated thecarcass of the trumpeter hornbill, she abandoned it
without eating it. In the 3rd case, although the alpha male
AL approached the meat possessor OS, AL did not show
any interest in the meat. In Gombe, among 14 episodesof bird predation by chimpanzees, 7 were performed by
the same individual—a juvenile female named Pom8.
Lastly, data across study sites show thatchimpanzees may not prey on birds with enough ability
to fly. Most of developmental stages of bird prey were25
17

Pan Africa News, 15(2), December, 2008
Table 1. Species, developmental stages of birds preyed upon by wild chimpanzees and age-sex of
predators in five chimpanzee study site.

chimpanzees of the Mahale Mountains. Primates 23:
364–384.

eggs, nestlings or injured adults, without either ability to
fly or resistance to predators.
6

Nishida T, Uehara S 1983. Natural diet of chimpanzees(Pan troglodytes schweinfurthii): Long-term record from
the Mahale Mountains, Tanzania. Afr Study Monogr 3:
109–130.

7

Takahata Y, Hasegawa T, Nishida T 1984. Chimpanzee
predation in the Mahale Mountains from August 1979 to
May 1982. Int J Primatol 5: 213–233.

8

Wrangham RW 1975. The Behavioural Ecology of
Chimpanzees in Gombe National Park, Tanzania. Ph.D.
thesis, Cambridge University.

9

Goodall J 1986. The Chimpanzees of Gombe: Patterns of
Behavior. Harvard University Press, Cambridge MA.

Acknowledgements. This study was financed by JSPS
Grant-in-Aid for Scientific Research (A1, #12375003 and
16255007 to Toshisada Nishida).

REFERENCES
1
2

Boesch C 1994. Cooperative hunting in wild chimpanzees.
Anim Behav 48: 653–667.
Hosaka K, Nishida T, Hamai M, Matsumoto-Oda A,
Uehara S 2001. Predation of mammals by the chimpanzeesof the Mahale Mountains, Tanzania. In: All Apes Great andSmall. Vol. 1: African Apes, Galdikas BMF, Erickson N,
Sheeran LK, Shapiro GL, Goodall J (eds), Plenum
Publisher, New York, pp. 107–130.

3

Newton-Fisher NE, Notman H, Reynolds V 2002. Huntingof mammalian prey by Budongo forest chimpanzees. Folia
Primatol 73: 281–283.

4

Watts DP, Mitani JC 2002. Hunting behavior ofchimpanzees at Ngogo, Kibale National Park, Uganda. Int J
Primatol 23: 1–28.

5

Kawanaka K 1982. Further studies on predation by

10 Kuroda S, Suzuki S, Nishihara T 1996. Preliminary reporton predatory behavior and meat sharing in tschego
chimpanzees (Pan troglodytes troglodytes) in the Ndoki
forest, northern Congo. Primates 37: 253–259.
11 Sugiyama Y, Koman J 1987. A preliminary list of
chimpanzees’ alimentation at Bossou, Guinea. Primates 28:
133–147.
12 Uehara S, Nishida T, Hamai M, Hasegawa T, Hayaki H,
Huffman MA, Kawanaka K, Kobayashi S, Mitani JC,

26
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Pan Africa News, 15(2), December, 2008

Takahata Y, Takasaki H, Tsukahara T 1992. Characteristics
of predation by the chimpanzees in the Mahale Mountains
National Park, Tanzania. In: Topics in Primatology, Vol. 1:
Human Origins, Nishida T, McGrew WC, Marler P,
Pickford M, de Waal FBM (eds), University of Tokyo Press,
Tokyo, pp. 143–158.
13 Huffman MA, Kalunde MS 1993. Tool-assisted predation
on a squirrel by a female chimpanzee in the Mahale
Mountains, Tanzania. Primates 34: 93–98.
14 van Perlo B 1995. Birds of Eastern Africa, Harper Collins
Publishers, London.

<NOTE>The Ebo forest: Four years of
preliminary research andconservation of the
Nigeria-Cameroon
chimpanzee (Pan troglodytesvellerosus)
Ekwoge E. Abwe1 and Bethan J. Morgan2
1. WCS-Ebo Forest Research Project, BP 3055 Messa,
Yaoundé, Cameroon

This report aims to introduce the four years of the
Ebo Forest Research Project (EFRP) which has theobjectives for research and conservation of the
Nigeria-Cameroon chimpanzees in the Ebo forest locatedin the south-western part of Cameroon.

METHODS
Research site
The Ebo forest extends over 2,000 km2 of closed
canopy primary forests in the biodiversity ‘hotspot’
between the Cross River in Nigeria and the Sanaga Riverin Cameroon (Fig. 1). The forest harbours eleven diurnal
primate species including the Nigeria-Cameroon
chimpanzee and a small population of gorillas of
uncertain taxonomic affinity2. Many small villages
surround the Ebo forest, which until the late 1950s andearly 1960s was inhabited mostly by the people of theBanen and Bassa tribes. Bushmeat hunting and trade are
the main economic activities in these villages, and thishas been facilitated by a network of logging roads
especially in the southern part of the forest. Following
results of preliminary surveys carried out in 2002, apermanent research station was set up in the Ebo forestby the Zoological Society of San Diego in April 2005
(Fig. 2).

2. Central Africa Program, Conservation and Research for
Endangered Species (CRES), Zoological Society of San
Diego, P.O. Box 3055, Messa, Yaoundé, Cameroon

INTRODUCTION
Cameroon harbours four great ape subspecies
separated by the Sanaga River, which is an importantbiogeographical barrier1. To the south are the sympatricwestern lowland gorilla (Gorilla gorilla gorilla) andcentral chimpanzee (Pan troglodytes troglodytes) and tonorth the Cross River gorilla (G. g. diehli) and therecently distinguished Nigeria-Cameroon chimpanzee
subspecies (P. t. vellerosus)1,2. The Nigeria-Cameroon
chimpanzee is the least studied of the four currently
recognised chimpanzee subspecies with the only current
long-term study being that in the Gashaka Gumti
National Park in Nigeria — a dry sub Saharan Guinea
zone3. Very little information is available on thissubspecies in the closed canopy primary forests of
Cameroon.

ProcedureLine transects and pre-existing human and elephanttrails were used to determine chimpanzee and gorilla
distribution and ranging patterns. Basic ecological data
was collected by indirect means, from beds and faecal
remains. Monthly observational censuses wereconducted along each transect and trail, noting
primate/mammal signs, feeding remains and human signs.
Faecal samples collected within the study area were
washed in 1 mm mesh metal sieves to distinguish thelarge seeds, other plant and animal matter ingested by theapes4.
Our outreach program targeted all age groups inmost of the villages around the Ebo forest. It consisted ofsporadic hunter training workshops, primate film shows,formal and informal education programs in schools andvillages.

27
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Pan Africa News, 15(2), December, 2008

Fig. 1. Location of the Ebo forest in
Cameroon.

Fig. 2. The Ebo forest area, research camps and surrounding
villages.

RESULTS
Bed structures
Of the 222 beds recorded along transects and trails
between January 2006 and December 2007, 24 wereground beds of which 6 were night beds (Fig. 3). The
Ebo chimpanzees constructed their beds mostly on steep
slopes and rugged terrain probably because of night
hunting by humans prevalent in the forest.

cracking was Coula nuts. The Ebo chimpanzees havenever been observed to crack open oil palm (Elaeis
guineensis) nuts like the Bossou chimpanzees in Guinea8.
Oil palms are common throughout the forest, and the red
soft outer layer is widely consumed by the chimpanzees.
So far, nut cracking sites have only been identified on thewestern side of the Ebo River. Four tool combinations
for cracking Coula nuts have been observed in the Ebo
forest: wooden anvil and wooden hammer, stone hammer

Termite fishing by chimpanzees
Discarded termite fishing tools were observed on
epigeal and subterranean nests across the Ebo forest (Fig.
4). As this activity has not been observed directly in the
Ebo forest, we are reluctant to speculate on how exactly
it is carried out. We have, however, noted two
combinations of instruments: stout long straight sticks
and flexible leaf midribs, similar to those found in theGoualougo Triangle in Congo5.
Nut cracking by chimpanzee
The Ebo chimpanzees were observed cracking
Coula edulis nuts in 20056, an activity hitherto thought tobe limited to communities west of the N’zo Sassandra
River in Côte d’Ivoire7. The target species for nut

Fig. 3. Fresh chimpanzee night-ground bed(photograph by EEA).

28
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Fig. 4. Termite fishing sticks on a subterranean
termite bed (photograph by EEA).

Fig. 5. Stone hammer and anvil (photograph by
EEA).

and wooden anvil, stone hammer and stone anvil (Fig. 5)
and stone hammer without anvil.

Sympatric gorillas
Gorillas were first sighted in the Ebo forest in
November 2002. This gorilla population isbiogeographically interesting as it occurs less than 100
km north of the Sanaga River, the geographical barrier
between the Western Lowland gorilla to the south andthe Cross River gorilla to the north2. The gorilla
population is sympatric with chimpanzees around the
Moupon area. From 2005 and 2007, ten groups of 31
gorilla beds were recorded 22 of which were tree beds
ranging between 0.5 m to 20 m in height. The groundbeds were constructed of vegetative matter on rocks orbare ground and in a couple of occasions, the gorillas
were observed to have slept on bare soil alongside
ground and tree beds.

Pan Africa News, 15(2), December, 2008
Outreach programs
The population of the 19 villages surrounding theEbo forest generate income largely from the commercial
trade in bushmeat. This coupled with local andcommercial logging activities are the main threats to theforest and fauna of Ebo. Hunters spend several days inthe forest and on their return to the village; they sell thebushmeat to intermediaries commonly called ‘buyam
sellams’ who in turn supply the cities. The ‘buyam
sellams’ come in hired taxis, motorcycles and sometimes
timber trucks based on appointments with the hunters.
Outreach programs were an integral part of the EFRP.
Four workshops for hunters from Ebo villages wereorganized at the Limbe Wildlife Centre for mammal
identification and comprehension of the national andinternational wildlife law. Three primate film shows
centred on the behavioural ecology of chimpanzees andgorillas were organized in three villages. Formal andinformal discussions on environmental issues wereregularly held in the villages and primary schools.
DISCUSSION
The long-term goal of the Ebo Forest Research
Project is to secure the population of chimpanzees,
gorillas and other large endangered mammals of the Ebo
forest. We are working on this through active researchcoupled with regular contacts with communities that use
the forest resources. Today we have two permanent
research stations (Bekob and Moupon) in the Ebo forest
and the third (Njuma) will be operational in January
2009. We are collaborating with the Government of
Cameroon and Wildlife Fund for Nature Coastal Forests
Programme for the gazettement of the Ebo forest into anational park. More international networking,collaboration and commitment are required to promote
the research and conservation of the endangered species
in the Ebo forest, including the Nigeria-Cameroon
chimpanzee.
Acknowledgements. Research and conservation in the Ebo
forest is funded by the Zoological Society for San Diego, theOffield Family Foundation, the United States Fish and Wildlife
Service (USFWS), Great Ape Conservation Fund, the USFWS
African Elephant Conservation Fund and the Margot Marsh
Biodiversity Foundation. In Cameroon, we work underauthorisation from the Government of Cameroon’s Ministry ofForestry and Wildlife and the Ministry of Scientific ResearchPan Africa News, 15(2), December, 2008
and Innovation. Our in-country partners include the Wildlife
Conservation Society, and WWF Coastal Forests Programme
Cameroon. We are grateful to the Ebo Forest Research Project
staff, chiefs and villages surrounding the Ebo Forest. Special
thanks to Professor Tetsuro Matsuzawa (Director of thePrimate Research Institute of Kyoto University) for inviting
EEA to the HOPE/SAGA symposium to introduce our research
and for reading and commenting on earlier drafts of thismanuscript.

REFERENCES
1

Gonder MK, Oates JF, Disotell TR, Forstner RMJ, Morales
JC, Melnick DJ 1997. A new West African chimpanzee
subspecies? Nature 388: 377.

2

Morgan BJ, Wild C, Ekobo A 2003. Newly discovered
gorilla population in the Ebo forest, Littoral Province,
Cameroon. Int J Primatol 24: 1129–1137.

3

Fowler A, Sommer V 2007. Subsistence technology of
Nigerian chimpanzees. Int J Primatol 28: 997–1023.

4

Tutin CEG, Fernandez M 1993. Fecal analysis as a methodof describing diets of apes: Examples from sympatricgorillas and chimpanzees at Lopé, Gabon. Tropics 2:
189–197.

5

Sanz C, Morgan D, Gulick S 2004. New insights into
chimpanzees, tools, and termites from the Congo Basin.
Amer Nat 164: 567–581.

6

Morgan BJ, Abwe EE 2006. Chimpanzees use stone
hammers in Cameroon. Curr Biol 16: R632–R633.

7

Boesch C, Marchesi P, Marchesi M, Fruth B, Joulian F
1994. Is nut cracking in wild chimpanzees a cultural
behaviour? J Hum Evol 26: 325–338.

8

Matsuzawa T 1994. Field experiments on the use of stone
tools in the wild. In: Chimpanzee Cultures, Wrangham RW,
McGrew WC, de Waal FMB, Heltne PG (eds), Harvard UP,
Cambridge, pp. 351–370.

<NOTE>

Coprophagy by thesemi-habituated chimpanzees
of Semliki, Uganda
C.L.R. Payne1, T.H. Webster1,2 and K.D. Hunt3
1. Leverhulme Centre for Human Evolutionary Studies,

University of Cambridge
2. Departments of Anthropology and Zoology, Miami

University
3. Department of Anthropology, Indiana University

29
17
INTRODUCTION
This report adds to the growing published literature
on coprophagy in wild chimpanzees. We present
observational and circumstantial evidence of thisbehavioural pattern, and test the utility of current
hypotheses for explaining coprophagy in wild
chimpanzees.
Eating one’s own feces (autocoprophagy) and eating
the feces of others (allocoprophagy) occurs in wild
populations throughout the animal kingdom1. Yet in apes,
coprophagy is commonly considered to be abnormal
behaviour in captive populations. It often appears in lists
of problematic behaviours2,3, and is thought to be a
response to boredom or stress4.
Yet wild chimpanzee populations also engage incoprophagy: Autocoprophagy has now been reported
from several long term study groups, including Assirik5,
Gombe6, Mahale7, Bossou8, and Fongoli (Bertolani et al.,unpublished data). Interspecific allocoprophagy has beenreported from Kibale, where chimpanzees consume
elephant dung9. The cause of this pattern remains
uncertain, but several hypotheses have been proposed.
(a) Boredom: Commonly cited as a factor inducing
coprophagy in captivity4, it has also been suggested
to explain wild gorilla coprophagy during long
periods of heavy rain when foraging activities are
reduced10.
(b) Insufficient roughage: Lack of “wadging” materials(such as fibrous leaves) in the diet of captive
chimpanzees may increase the frequency of
coprophagy11.
(c) Provision of essential nutrients: Some captive
groups of largely herbivorous gorillas may engage
in coprophagy in order to gain Vitamin B12 which
is present only in animal matter12.
(d) Food scarcity: At Gombe, coprophagy occurred
during a period of fruit scarcity, after an unusually
dry season6. The pressures of foraging in afood-scarce environment may increase coprophagy.
(e) Reingesting hard seeds: Coprophagy may be
adaptive when groups feed on fruits with seeds thatpass through the gut whole, but then can be split
open and reingested after coprophagy. Free-ranging
chimpanzees in the Republic of Congo regularly
engaged in coprophagy when feeding on Dialium

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(f)

Pan Africa News, 15(2), December, 2008
seeds, which were found both whole andfragmented in feces13. Chimpanzees at Assirik
extract and eat baobab (Adansonia digitata) seeds
from their feces5. Allocoprophagy in Kibale ispositively correlated with the presence of hard
seeds in the elephant dung9. In Mahale, anindividual showed coprophagy after eating Saba
florida7, a species with hard seeds that are ingested
whole.
Culture: Nash et al.3 presented evidence suggestingthat coprophagy may be a socially learned
behaviour in captive chimpanzee groups.

METHODSThe Toro-Semliki Reserve is in western Uganda,
near the eastern edge of the Great Rift Valley (0°50’ to
1°05’ N, 30°20’ to 30°35’E), and supports several
chimpanzee communities14. Habituation efforts havebeen ongoing since 199615, and progress is being made.
We made observations and collected fecal samples
opportunistically from May–November 2008. Feces were
washed and sieved through a 1 mm mesh, and thecontents recorded.

RESULTSWe saw five episodes of autocoprophagy during thestudy period (See details in Table 1). No allocoprophagy
was observed directly, but circumstantial evidence ofcoprophagy comes from one case of discarded(presumably spat-out) seeds (Table 1), and from seedfragments in eight fecal samples (Table 2).
At least three individuals, including adult andjuvenile males, showed coprophagy (Table 1).
Individuals were observed to defecate directly into theirown hand and raise the feces to their mouth. They thenmanipulated the feces in their mouths using their lips,
spitting out seeds and indeterminate fecal matter. These
cases occurred when groups ate low-quality food items,
such as bark, pith and seeds (Table 1). Case 6 occurred
when at least one individual was in poor health, asevidenced by vomit found on leaves, although noevidence of disease or infection was found at any othertime during the study.
The presence of seed fragments in feces may be
indirect evidence of coprophagy, because the apes were
never seen or heard to crush the hard Saba florida seeds
when eating the fruits whole. In every case in which
coprophagy was observed, we could hear the sounds ofTable 1. Observed cases of coprophagy by Semliki chimpanzees.

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Pan Africa News, 15(2), December, 2008
Table 2. Chimpanzee fecal samples containing Saba florida seed fragments.

Fig. 1. Indirect evidence of coprophagy from fecal analysis: S. florida consumption.

seeds being crunched and cracked. We presumed these to
be Saba, since these seeds were predominant and were
found more consistently in fecal samples than any otherseeds during the study period. We found crushed seeds insamples in the latter half of the Saba fruiting season (Fig.
1), shortly before Saba seeds disappeared from feces
altogether.
The decline in the presence of whole seeds andincrease in the presence of fragments in feces between
Weeks 1–8 and Weeks 9–16 (Fig. 1) differs significantly
from chance (X2 (df=1) = 10.7, N=78, p<0.01).

DISCUSSION
The hypotheses (a)–(b) cannot explain coprophagyby the Semliki chimpanzees. (a): Unlike barren captive

environments, the habitat at Semliki reserve is rich andstimulating; the chimpanzees engage in various kinds ofextractive foraging. (b): Fibrous materials often featurein the diet of the Semliki chimpanzees14, and wereconsumed during every case of observed coprophagy.
(c): Insects occurred regularly in fecal samples at
Semliki during the study period, so coprophagy need notsupply Vitamin B12.
Instead, hypotheses (d) and (e) seem likely variables
which may interact to cause the chimpanzees of Semliki
to engage in coprophagy. Given the significant
relationship between the presence of crushed seeds infeces and declining Saba availability, we propose atentative hypothesis that coprophagy in Semliki
chimpanzees is caused by the pressure to exploit added

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nutrients from reingesting Saba seeds when availabilityof the fruit is in decline. There is a possibility that seedfragments in feces may not reflect coprophagy but
instead the occasional crunching of the seeds during
Saba fruit consumption that is never seen at Semliki, butobservedat
Mahale(Nakamura,personal
communication). Further observational data are needed
to clarify this.
The lack of coprophagy during times when Saba
was consumed in abundance, and the presence of many
whole, non-reingested seeds in feces in the early half ofthe fruiting season, suggests that coprophagy may be a
fallback or seasonal strategy for Semliki chimpanzees. It
may be a response to a decreasing resource, as a way ofmore efficiently exploiting the nutrients available fromthis high quality food item. It may also be specific toindividuals who find it harder to acquire high quality
foods (Table 1). However, further behavioural data isneeded to test this, particularly since coprophagy was
only observed in adult and juvenile males and these
individuals are better habituated to human observation
than female chimpanzees at Semliki.
Hypothesis (f) cannot be assessed from the data
gathered thus far. As habituation efforts progress, it ishoped that more information on unusual behavioural
patterns in this population will come to light.
Acknowledgements. We thank the National Science Foundation,
University of Indiana, Rebecca Jeanne Andrew Memorial Fund,and Bedford and Durham Funds of King’s College, Cambridgefor financial support; WC McGrew and LF Marchant for help
with data collection and analyses; and the rangers of the
Ugandan Wildlife Authority and the staff of Semliki
Chimpanzee Research Camp, for field assistance.

Pan Africa News, 15(2), December, 2008
TL, Mitchell G (eds), Van Nostrand, New York.
5

Baldwin PJ 1979. The Natural History of the Chimpanzee(Pan troglodytes verus) in Mt Assirik, Senegal. PhD thesis,
University of Stirling.

6

Goodall J 1986. The Chimpanzees of Gombe: Patterns of
Behavior. Belknap Press, Cambridge MA.

7

Uehara S 1979. The chimpanzees of Kasoge K group [inJapanese]. Monkey 170: 16–25 (Cited in Krief et al. 2004)

8

Nishida T, Kano T, Goodall J, McGrew WC, Nakamura M
1999. Ethogram and ethnography of Mahale chimpanzees.
Anthropol Sci 107: 141–188.

9

Krief S, Huffman MA, Sévenet T, Guillot J, Bories C,
Hladik CM, Wrangham RW 2005. Noninvasive monitoringof the health of Pan troglodytes schweinfurthii in the
Kibale National Park, Uganda. Int J Primatol 26: 467–490.

10 Harcourt AH, Stewart KJ 1978. Coprophagy in wild gorilla.
E Afr Wildl J 16: 223–225.
11 Fritz J, Maki S, Nash LT, Martin T, Matevia M 1992. The
relationship between forage material and levels of
coprophagy in captive chimpanzees (Pan troglodytes). Zoo
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Pan Africa News, Vol. 15, No.2
Published in December, 2008
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