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/

DECEMBER 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 Mts. Chimpanzee Research Project
Eiji Inoue, Kyoto University, Japan
Instructions for 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 of conservation and research regarding chimpanzees (Pan troglodytes)
and bilias (Pan paniscus). Contributors are requested to writein English and the papers except forums, reviews and essays
should usually be 1,500 words or less. Articles, notes and reviews 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 andfigures should be formatted as JPEG or GIF files and sent separately by e-mail attachments. Authors could include audiovisual data to enhance their papers, although they will be includedin the online version only. Sound or video files should be sent
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PAN is published twice a year in June and December. Deadline for manuscripts is one month before publication (i.e. theends of May and November).

<NEWS>GRASP in Paris 2012
Tetsuro Matsuzawa1,2
1 Primate Research Institute, Kyoto University, Japan
2 President of International Primatological Society(E-mail: matsuzaw@pri.kyoto-u.ac.jp)

GRASP, which stands for the Great Apes Survival
Partnership, is a joint program of UNESCO and UNEP.
The 1st GRASP council meeting was held in Kinshasa,
Congo, in 2005. The discussion at the time resulted in theVOL. 19, NO. 2

Contents<NEWS>GRASP in Paris 2012
Tetsuro Matsuzawa			
15<NEWS>
Iyondji Community Bonobo Reserve: A Recently
Established Reserve in the Democratic Republic ofCongo
Tetsuya Sakamaki, Phila Kasalevo, Mathieu
Bolaa Bokamba & Lingomo Bongoli		
16<NOTE>
Death of the Oldest Female at Mahale and Some Notes
about Longevity of Wild Chimpanzees
Michio Nakamura & Hitonaru Nishie		
19<NOTE>
Wild Chimpanzees at Mahale are not Manually
Lateralised for Throwing
Toshisada Nishida, William C. McGrew &Linda F. Marchant 			21<NOTE>
Playful Drumming by Immature Wild Chimpanzees at
Mahale: Do They Enjoy Making Sounds?
Takahisa Matsusaka			
23<NOTE>
Comparison of the Longevity of Chimpanzee Beds
between Two Areas in the Mahale Mountains National
Park, Tanzania
Koichiro Zamma & Masayuke Makelele
25<BOOK INFO>
Wild Cultures					
28Kinshasa Declaration on Great Apes. The 2nd meeting of
the GRASP council was held in November 6–8th, 2012, inthe headquarters of UNESCO, Paris, France. The author
is a member to the Scientific Commission (SC) of GRASP.
The SC consists of experts on wild great apes and acts toprovide scientific advice to GRASP.
The GRASP 2012 council meeting in Paris aimed todefine and prioritise actions for the coming years. The
new GRASP coordinator, Douglas Cress, took the leadat this international gathering, which included predominantly representatives of range countries, donor countries,
NGOs and UN-related organizations. I estimate that about

15

16

Pan Africa News, 19(2), December 2012

GRASP can, for instance, help to influence policy-makers, to promote green
economy strategies, and to improve law
enforcement coordination worldwide
to the benefit of great apes. This levelof effort is essential if we are to resolve
conservation challenges lying beyond
national borders, producing a real impact on conservation through policymaking whilst balancing conservationand economic development.
I am writing this short note in
Bossou, Guinea, West Africa, where
the Kyoto University team has been
carrying out a longitudinal study ofchimpanzees in Bossou and Nimba.
Among other tool uses and remarkablebehaviors, Bossou chimpanzees aremost well-known for their use of a pairof mobile stones to crack open oil palm
nuts. However, in recent years, theFigure 1. The 2nd GRASP Council was held 6–8 November 2012 at UNESCO Bossou chimpanzees have been declinhead­quarters in Paris.
ing in numbers, with only 12 individu150 people gathered for this 3-day meeting.
als remaining today. If this community were to become
There was a lot of exchange of information about the extinct, its unique culture will disappear with it. Mountcurrent situation of great apes and major threats to their Nimba, 4 km away from Bossou, is the only natural World
survival including habitat loss, poaching and the bush- Heritage Site (WHS) in Guinea; however, this mountainmeat trade, as well as the illegal international trade and the range, which is rich in high grade iron-ore, continuously
risk of disease transmission. The core of the discussions faces the threat of mining. We have, for many years now,and output documents are available on the GRASP web- been taking the initiative to plant trees between Bossou
site. Please visit the website (http://www.un-grasp.org/) for and Nimba to connect the two areas while also encouran update on GRASP activities. This report simply aims aging natural regeneration; this project is known as theto publicize that the second GRASP council meeting was Green Corridor Project.
held in Paris to revitalize conservation efforts for wild
Science in the field should accompany in situ consergreat apes across their range.
vation efforts, just like science in the laboratory should
There are many ongoing activities at different lev- aim to support welfare of captive animals. In various
els aimed at promoting conservation of great apes in the places, field scientists are making their own grass-roots
wild. In this context, it might be helpful to mention the efforts. I hope that activities at all levels will complement
different roles of three organizations: the International one another to improve the conservation of great apes andPrimatological Society (IPS), IUCN, and GRASP.
ensure their long-term survival in the wild.
IPS is the international academic society of primatologists. It holds biennial meetings. IPS 2010 was held in
Kyoto, Japan, IPS 2012 was just held in Cancun, Mexico,and IPS 2014 will be held in Hanoi in Vietnam. The IPS<NEWS>
comprises scientists across various disciplines who study
non-human primates (NHPs). The IPS plays a key role inthe scientific approach to conservation of NHPs, because
the majority is either endangered or threatened. It is very
important to acknowledge that there exists a wide variety
of primate species, other than just great apes. IPS has avice-president for conservation.
The IUCN species survival commission (SSC) also
Tetsuya Sakamaki1,2, Phila
unites scientists, taxon experts whose role is, among othKasalevo3, Mathieu Bolaa
ers, to evaluate the conservation status of species andBokamba4 & Lingomo Bongoli5,6
regularly update the IUCN Red List of threatened spe1 Primate Research Institute of Kyoto University, Japancies. The IUCN SSC includes a Primate Specialist Group
2 Wamba Committee for Bonobo Research(PSG). The PSG has a Section on Great Apes (SGA). The
3 African Wildlife Foundation
4 Ecosystem Restorate Associate (ERA-Congo)
IUCN-PSG-SGA consequently focuses on the conserva5 Forêt des Bonobos
tion of great apes from a scientific viewpoint.
6 Centre de Recherche en Ecologie et Foresterie, D. R. Congo(E-mail: sakamaki.tetsuya.8v@kyoto-u.ac.jp)
GRASP differs from IPS and IUCN-SSC in its ability
for advocacy. As part of the United Nations organization,

Iyondji Community Bonobo
Reserve: A Recently Established
Reserve in the Democratic
Republic of Congo

Pan Africa News, 19(2), December 2012
INTRODUCTION

The Iyondji Community Bonobo Reserve (ICBR), established by the Democratic Republic of the Congo (DRC)
Ministry of Environment in April 2012, encompasses approximately 1,100 km² of tropical moist forest that harbors
bonobos (bilia; Pan paniscus). The ICBR is adjacent to thesoutheast border of the Luo Scientific Reserve (481 km²).
Wamba, a long-term study site of wild bonobos, is situated in the northern section of the Luo Scientific Reserve1(Figure 1). The Kokolopori Bonobo Reserve is east of the
ICBR. Creation of the ICBR will benefit the conservationof bonobo habitat because it expands the protected area.
The present report describes the history of the area before
the ICBR was gazetted, efforts to habituate the bonobos,
and current management issues and implications for the
future.

BACKGROUND

Most Iyondji villagers live in settlements that extend
along established roads in the northern section of the Luo(Maringa) River. Crop fields and secondary forests stretch
for 1 to 2 km on both sides of the villages. Primary forest extends along the southern section of the river where
temporary hunting, fishing and gathering camps andsmall hamlets with cassava fields lie2 (Figure 1). The local Iyondji communities (Yohala and Yokali villages)
wanted to establish a community reserve in the southern
section of the Luo River, and in 2007, they requested help
from the African Wildlife Foundation, an international
non-government organization (NGO), and the Japanese
research team that had established the Wamba Committee
for Bonobo Research, a long-term research project in andaround Wamba. At the time, we conducted preliminary
surveys and continued to train and support the community in searching for local bonobo group, and a local NGO,Forêt des Bonobos, took the initiative and carried out the17
project with the villagers. Between 2010 and 2011, withsupport from the United States Fish and Wildlife Service,efforts to gazette a new community reserve were made inearnest.

HABITUATION OF BONOBOSIn July 2010, habituation of the bonobos was begun
on a daily basis in preparation for potential tourism or
scientific research. The habituation procedure involved
identifying unit-groups (or communities) and their ranges
by finding footprints and food remnants, making andmaintaining observation trails in the ranges, following
one or two target groups on a daily basis and identifying
individuals in the bonobo groups. We established a research camp at a hamlet along the Bembongo River in thenorthern part of the proposed protected area (0° 8′ 20″ N,
22° 44′ 37″ E; Figure 1). The Bembongo camp was 2–3 hfrom an Iyondji village (outside of the proposed protectedarea) travelling in a small boat and on foot.
We found two different groups of bonobos aroundthe camp in the first 2 months. We focused our efforts on
the Bembongo Group and followed them on a daily basis.
Table 1 shows the number of days the group was observed
and the direct observation times during the project. The
camp was situated in the northern sector of the Bembongo
Group’s range. It was possible to follow the bonobos
nearly every day when they were near the camp, but thiswas difficult when they ranged further south because theirsouthern range was 2 h or more from the camp and we
had not made observation trails further south.
We identified the Ite Group, a unit-group that rangedin the west adjacent area and the Bohondohondo Group,
which ranged in the east adjacent area. Moreover, wefound evidence of other bonobo groups in the southernarea of the reserve.
The ICBR bonobo population borders the Wamba

Figure 1. Map of the Iyondji Community Bonobo Reserve and the Luo Scientific Reserve.

Pan Africa News, 19(2), December 201218

Table 1. Number of research days, direct observations and observation time of
the Bembongo group.

2010
July
AugustSeptember
OctoberNovember
December
2011
January
February
March
April
May
June
July
AugustSeptember
OctoberNovemberNumber of
research days

Number ofobservation days

Observation time (h)

17
8
12
0
26
26

3
0
0
–
4
13

0.3
0
0
–
19.32
33.18

23
28
31
27
31
30
31
30
27
30
25

12
16
23
11
19
15
12
12
7
16
12

45.38
62.62
66.9
24.28
84.63
32.13
27.58
27.57
23
30.52
44.63

bonobo population. Behavioural and ecological studies of the Wamba group have been reported previously1.
Given the proximity of the two groups, their habitats areunlikely to differ significantly; however, further surveys
are necessary to assess the two environments. Individuals
in the ICBR and Wamba groups cannot transfer between
groups because they cannot cross the Luo River, which
separates the two populations. Thus, the behavioural diversity or “culture” of the ICBR bonobos is an interesting
topic for future study3.

CURRENT PROBLEMS AND THE FUTURE

The local Bongando people (a Bantu ethnic group) areclosely tied to their natural environment2. Although theirmost important crop is cassava, the Bongando acquire
most of their animal protein from the forest where theyhunt and gather various species of mammals, birds, reptiles, fish, insects and plants. They keep livestock such as
goats, pigs, chicken and ducks, which are consumed in rituals and on ceremonial occasions. Effective and sustainable management of the ICBR requires the developmentof alternative sustainable livelihoods while controlling theuse of resources in the forest.
People living in the forest present additional difficulties. One objective of the 2010–2011 Iyondji project was toestablish a biannual monitoring of large mammals and human activity in the proposed protected area. We originallyplanned to make 10 line transects (each 5 km) in the areafor biannual monitoring and regular patrol. However, ourmonitoring walks were able to cover only four transects
in the northern area because the people who lived in scattered hamlets in the middle and southern areas of the proposed protected area were not in favour of a new reserveand obstructed our work. These were not Iyondji people,

but members of other populations in the adjacent administrative districts. Although the southern boundary of the
ICBR was within the Iyondji jurisdiction, the Iyondji people only inhabited the forest in the northern sector of the
ICBR.
Temporary hunting and gathering camps and small
hamlets with cassava fields were present within the ICBR.
Approximately 10 people lived in a hamlet (Lingomo etal., unpublished data). According to the villagers, people
lived in the forest permanently, and some of their children
had never been to the villages along the established roads
where there were schools and dispensaries. This situation
was, in part, the result of the DRC civil war in the 1990s2.
Prior to the civil war, villagers hunted primarily for theirown consumption. However, during the war the trucking
and shipping transportation network collapsed, and thevillagers were left with no means of selling cash crops
such as coffee beans. As a result, the sale of bush meat
became the primary way of earning cash.
Unlike the people from adjacent districts, the local Iyondji communities agreed to manage the ICBR,
and they regularly monitor and patrol their forest. It is
important to note that the Iyondji people were provided
with opportunities for alternative livelihoods that allowed
them to decrease the hunting of wild animals and habitat
destruction through slash-and-burn agriculture. Continual
support for the Iyondji and other adjacent communities
may be necessary to resolve conf licts with the ICBR.
Furthermore, it is necessary to develop innovative ways tocreate revenue through the forest and its bonobos, such as
tourism and scientific research.
At present, it is not realistic for the local communities to forego the use of resources in the forest. However,
local community activities must adhere to regulations ofPan Africa News, 19(2), December 2012
the reserve, and regular monitoring of animals and human activity is important and necessary. Ideally, the local communities will take the initiative in carrying out
regular monitoring and patrolling of their forests. Our
ICBR project is a model case of collaboration among local
communities, conservation NGOs and scientific researchers. At this stage of the ICBR project, the collaboration is
expected to continue to help solve current problems andfacilitate development of the management plan.

ACKNOWLEDGEMENTS

Invaluable support and logistical assistance were provided
by Jef Dupain, Charly Facheux, Lillian Nsimba and other members of the Africa Wildlife Foundation (AWF); the Centre deRecherche en Ecologie et Foresterie (CREF) of the Democratic
Republic of Congo; T. Furuichi, D. Kimura and other membersof the Wamba Committee for Bonobo Research; members of
the local NGO, Forêt des Bonobos, and the local Iyondji communities. Research permission was arranged by the CREF
and the Ministry of Scientific Research of the Democratic
Republic of Congo. I thank K. Hosaka and T. Furuichi for useful comments on the original manuscript. The present project
was financially supported by the US Fish and Wildlife Service
Assistance Award (96200-0-G017 to AWF), Grants-in-Aid
for Scientific Research of the Japan Society for the Promotionof Science (JSPS) (2225507 to T. Furuichi), the Environment
Research and Technology Development Fund of the Ministry of
the Environment, Japan (D-1007 to T. Furuichi), and the JSPS
Primate Origins of Human Evolution (HOPE) project (AS-22027, ITP-23-006 to T. Sakamaki).

19
on 5th April 2012. As it is rare to observe the deaths ofwild chimpanzees directly, or to find the dead bodies, weusually assume individuals who have not been observed
for more than 3 consecutive months in Mahale to be
deceased.
Calliope was estimated to be 52 years old in 2012,
which is the oldest age at death so far recorded at Mahale.
She was first identified in 1973 as a young adult. The
starting age of adulthood in females is 13 years old.
Therefore, she would have been at least 13 years old atthat time. This gave the estimation that she was born in
1960, but her exact birth-year could be even earlier.
According to Nishida1, Calliope had been very shy tohuman observers until the 1990s. Although she later became tolerant of observations from a certain distance, she
continued to be more timid than the other females. She
seemed reluctant to get too close to humans, especiallywhen she had a small baby. Throughout her life, Calliope
gave birth to a minimum of 5 offspring (i.e., 3 females
and 2 males)2. Except for 1 female offspring who died at
3 years old, the other 4 were weaned and reached the ageof puberty. Thus, it can be said that she was a successful
mother. After she gave birth to her last offspring in 1997,
she did not give birth again throughout the last 15 yearsof her life. Excluding the 5 years during which she wasnursing her last baby, she enjoyed her remaining postreproductive life for 10 years.

REFERENCES

1. Furuichi T, Idani G, Ihobe H, Hashimoto C, Tashiro Y,
Sakamaki T, Mulavwa MN, Yangozene K, Kuroda S
2012. Long-term studies on wild bonobos at Wamba, Luo
Scientific Reserve, D. R. Congo: Towards the understanding of female life history in a male-philopatric species. In:
Long-Term Field Studies of Primates. Kappeler PM, Watts
DP (eds), Springer, Heidelberg, pp. 413–433.
2. Lingomo B, Kimura D 2009. Taboo of eating bonobo
among the Bongando people in the Wamba region,
Democratic Republic of Congo. Afr Stud Monogr
30:209–225.
3. Hohmann G, Fruth B 2003. Culture in bonobos? Betweenspecies and within-species variation in behavior. Curr
Anthropol 44:563–571.

<NOTE>

Death of the Oldest Female at
Mahale and Some Notes about
Longevity of Wild Chimpanzees
Michio Nakamura & Hitonaru
NishieFigure 1. Calliope in 2010 (at estimated age of 50 yearsold). Although her hair was white, she did not look old.

Kyoto University, Japan(E-mail: nakamura@wrc.kyoto-u.ac.jp)

Death of Calliope, the Oldest Femaleat Mahale

Calliope (Figure 1), an old, female chimpanzee at
Mahale, is assumed to have died this past year (i.e., 2012).
Her death is assumed because it has not been confirmed,
but she has not been witnessed since the final observation

Calliope’s Final Days of Life

Calliope was often observed traveling with her last
daughter, Carmen, until Carmen emigrated from the M
group in July 2011. Calliope also groomed often with
her two mature sons, Carter and Cadmus, when she met
them. After the emigration of Carmen, Calliope often

20
ranged together with Wakusi, an older female who is estimated to be one year younger than Calliope. As the oldest
duet, they usually kept a moderate distance from noisy
males, even though they both had mature sons. However,in large, they ranged alongside the other chimpanzees.
We sometimes observed that they would feed apart fromthe others, whose calls could be heard from a distance.
Some moments later, they would travel in the directionof the calls in tandem. When they caught up with a groupof males, the males would often groom the two older females. The females, however, seldom groomed the males
in return, as if they were enjoying respects for the aged.
Calliope might have suffered from presbyopia. For instance, when she groomed others, she straightened up her
upper body so that her eyes were at a distance from thegroomee’s body surface.
When MN observed Calliope on 15th December
2011, she was, again, travelling with Wakusi. When thesetwo, and a mother-infant pair, were resting under a bush,there came her fully adult son, Carter. Then, Calliope
and Carter started to play with each other. They tickled,slapped, and mouthed each other’s body and chased eachother in a circle. Both of them showed a play face and playpanted. A 51-year-old mother and a 26-year-old adult son
were playing like two juveniles! The play lasted for 23
minutes.
HN was the last researcher to see Calliope on 4th
March 2012, when he was following an adult male who
was in consort with a young estrous female. On thisday, again, Calliope and Wakusi were together. The twojoined the party that HN was following. Although theobservation time for Calliope was short, HN did not notice anything peculiar about her health or physical condition. After that day, she was not observed again for sometime. This is not uncommon in March–April, however, as
Mahale chimpanzees are usually dispersed in small parties throughout this season. On 5th April 2012, a research
assistant observed Calliope with 10 other chimpanzees.
This was the last day that she was observed.

Longevity of Wild Chimpanzees

Establishing the longevity, or the maximum life­
span, of wild chimpanzees is difficult to ascertain.
Chimpanzees typically have a long lifespan, and theirages are often estimated. Let us compare, here, the 52
years of life of Calliope (the longest documented lifespanat Mahale) with other wild chimpanzee populations. At
Gombe, the oldest estimated age at death was 53 years
for the famous chimpanzee, Flo3. At Bossou, the oldest
female, Kai, died at an estimated 53 years of age4. At Taï,a female, Chanel, died when she was 46 years old (calculated from Appendix Table A.1. in ref. 5). These ages are
all within the estimated range of the maximal lifespan potential of captive chimpanzees (48 ± 5 years; taken from aliterature survey covering over 100 zoos worldwide; citedin ref. 6).
We have to be careful about the specific age at deathfor chimpanzee individuals in the wild because they are
all estimates. For example, Flo’s age at death was firstestimated to be 43 years old by Goodall7, but she was estimated to be 10 years older in the later literature3. ThisPan Africa News, 19(2), December 2012
may be because Flo was already old, in her 30s or 40s,
when she was first identified in 1962. Estimating the ageof older individuals is particularly difficult because individuals differ in their appearance. Like humans, some
chimpanzees look younger and others look older than they
really are. Accordingly, we see, with a bit of suspect, thata 55-year-old mother gave birth to a child at Kibale8. Thismeans that, if the estimation of age is correct, her longevity will be much longer than that or previously recorded
individuals. Research at Kibale was initiated in 19839,
which means this female was in her 30s at that time. Thus,
there may the possibility that her age was overestimated.
For the estimated age of Calliope, we expect a small
proportion of error because she was estimated to be 13
years old when she was first identified. It is possible that
one would estimate a 33-year-old female to be 43 years
old, but it is not likely that one estimates a 3-year-old female to be 13 years old. From our experience, estimation
error of a 13-year-old female can be within 3 years or so,thus we can more confidently say that Calliope was between 49 and 55 years old at the time of her death.
Finally, we should mention some aged individuals
who are still alive. At the time of writing, there wereseveral older chimpanzees at Mahale, such as Wakusi(female, 51), Gwekulo (female, 50), Fatuma (female, 49)
and Kalunde (male, 49) (numbers in parentheses are theirestimated ages in 2012, which are equally accurate as, oreven more accurate than, that of Calliope because these
individuals were in their young adulthood or adolescencewhen identified). Some of these individuals now lookold in their appearance and show the typical behavioral
characteristics of old age, such as slow traveling speed
and a relative decrease in activity. However, at present,
they all appear to be in good health. Similarly, at Bossou,
4 females and 1 male older than 50 years were still alivein 2012 (Ohashi G, personal communication). Looking
at these figures, although the sample size is still small, it
seems that more females experience old age than males
at both sites. This assumption concurs with the fact thatmales, on average, show higher mortality than females10.
We hope that these older chimpanzees will live longer, well beyond the current records of longevity in wild
chimpanzees. Then, our current knowledge about the longevity of wild chimpanzees would be revised.

Acknowledgements

We thank COSTECH, TAWIRI, and TANAPA for theirpermissions to conduct long-term research at Mahale; Prof.
Y. Sugiyama for constructive comments on the manuscript;
Mr. G. Ohashi for updated information on Bossou chimpanzees; and researchers of the Mahale Mountains Chimpanzee
Research Project for their contribution to the accumulationof demography data of Mahale chimpanzees. MN’s fieldstudy was supported by Kakenhi (#19255008, #19107007, and#21770262) and HN’s field study was supported by the JSPS
Institutional Program for Young Researcher Overseas Visits.

References

1. Nishida T 2010. Introduction to Mahale Chimps: #16
Calliope. Mahale Chimpun 16:6, in Japanese.
2. Nakamura M, Nishida T 2012. Long-term field studies of chimpanzees at Mahale Mountains National Park,
Tanzania. In: Long-Term Field Studies of Primates.
Kappeler PM, Watts DP (eds), Springer, Heidelberg, pp.

Pan Africa News, 19(2), December 2012
339–356.
3. Williams JM, Lonsdorf EV, Wilson ML, SchumacherStankey J, Goodall J, Pusey AE, 2008. Cause of deathin the Kasekela chimpanzees of Gombe National Park,
Tanzania. Am J Primatol 70:766–777.
4. Sugiyama Y and Fujita S 2010. The demography and reproductive parameters of Bossou chimpanzees. In: The
Chimpanzees of Bossou and Nimba. Matsuzawa T, Humle
T, Sugiyama Y (eds), Springer, Tokyo, pp. 23–34.
5. Boesch C, Boesch-Achermann H 2000. The Chimpanzeesof the Taï Forest. Oxford University Press, Oxford.
6. Cutler RG 1984. Carotenoids and retinol: Their possible
importance in determining longevity of primate species.
Proc Natl Acad Sci USA 81:7627–7631.
7. Goodall J 1986. The Chimpanzees of Gombe. Harvard
University Press, Cambridge, MA.
8. Emery Thompson M, Jones JH, Pusey AE, BrewerMarsden S, Goodall J, Marsden D, Matsuzawa T, Nishida T,
Reynolds V, Sugiyama Y, Wrangham RW 2007. Aging andfertility patterns in wild chimpanzees provide insights into
the evolution of menopause. Curr Biol 17:2150–2156.
9. Wrangham RW, Clark AP, Isabirye-Basuta G, 1992.
Female social relationships and social organization ofKibale Forest chimpanzees. In: Topics in Primatology,
Vol.1. Nishida T, McGrew WC, Marler P, Pickford M, deWaal FBM (eds), University of Tokyo Press, Tokyo, pp.
81–98.
10. Hill K, Boesch C, Goodall J, Pusey A, Williams J,
Wrangham R 2001. Mortality rates among wild chimpanzees. J Hum Evol 40:437–450.

<NOTE>

Wild Chimpanzees at Mahale
are not Manually Lateralised forThrowing
Toshisada Nishida1, William C.
McGrew2 & Linda F. Marchant3
1 Deceased
2 Div. of Biological Anthropology, Dept. of Archaeology &Anthropology, Univ. of Cambridge, UK
3 Dept. of Anthropology, Miami University (Ohio), USA(E-mail: wcm21@cam.ac.uk)

INTRODUCTION

The importance of throwing has figured consistently
and prominently in scenarios about the evolutionary origins of human behaviour1,2 The utility of imparting force
to airborne projectile weapons, launched ballistically, isobvious, whether to deter, punish or subdue predators,
prey, or competitors. Other functions of throwing are lessobvious but could be equally useful, such as bringing
down hanging fruit, gaining others’ attention, shatteringlarge objects into fragments, etc. Scenarios have stressed
a number of important variables, such as posture (especially bipedality), sensorimotor skill (especially hand-eye
coordination), cerebral asymmetry, etc. Calvin1 linked allof these to the evolutionary origins of language. However,few of these ideas have been tested empirically.
One way to tackle evolutionary aspects of human
throwing is to look at throwing performance in our nearest living relations. Although throwing has been known
in both wild 3 and captive 4 chimpanzees for almost a
century, quantitative studies are few; often they are sub-21
sumed in broader studies of manual behaviour, e.g. ref.
5. Furthermore, studies often do not distinguish between
aimed (targeted) throwing versus unaimed (perhaps better termed hurling) throwing, underarm versus overarm
delivery, or one-handed versus two-handed throwing.
Several studies of manual laterality in throwing by
captive chimpanzees have been published, and all havereported population-level, right-sided bias6–8. Others haveincorporated these reported findings into comprehensive
accounts of the origin of handedness, e.g. ref. 9
We can find no published quantitative data on manual
laterality in throwing from any non-human primate species in nature, much less from chimpanzees. One obvious
reason for this absence is that, unlike in the artificial conditions of captivity, wild primates throw only rarely. Here
we report such a dataset, collected ethologically from a
population of chimpanzees in nature, over an extended
period, in an effort to balance the picture.

METHODS

The subjects were the eastern chimpanzees (Pantroglodytes schweinfurthii) of M group in the Mahale
Mountains of western Tanzania10,11. This population has
been studied since 1965, making it the second-oldest fieldstudy of chimpanzees; the apes are fully habituated toclose-range behavioural observation. The Mahale ethogram is well-known and described; Nishida et al.12 listed
10 kinds of throwing.
We recorded all observed occurrences of throwing, using ad lib. sampling, that is, regardless of whether
behaviour was being scan-sampled or focal-subjectsampled13. We interrupted normal protocols, in order tonote the identity of the thrower, target, etc. Nishida recorded his data from 1999–2004, when he mostly followed
males, hence the shortage of data on females. McGrew
and Marchant recorded their data in 1996. Here we report
data only on unimanual versus bimanual data, and on thehand(s) involved. Each throw is considered an independent
data-point, as the chimpanzees always repositioned themselves between throws, usually to pick up another object.

RESULTSWe recorded 556 throws by 16 individuals (Table 1);
the median number of throws per subject was 33 (range:
9–72). Of these, 63 (11%) were done two-handed, usuallyin agonistic charging displays; almost half of these weredone by two high-ranking adult males (DE, n = 17; FN, n= 13). These appeared to be unaimed and perforce weredone bipedally; they are considered no further here.
One-handed throwing was unlateralized. Only 4 (BB,CT, IV, OR) of the 16 individuals showed statisticallysignificant (Binomial test, two-tailed, p < 0.05) laterality;
three were biased to the right versus one to the left The
remaining 12 subjects showed ambilateral performance.
Overall, nine individuals showed (non-significantly) more
right-sided throws, six showed left-sided throws (a nonsignificant difference, n = 15, x = 6, p = 0.60), and one
was tied. Descriptively overall, of one-handed throws, 226(46%) were done with the left hand versus 267 (54%) withthe right hand.

Pan Africa News, 19(2), December 2012

22

Table 1. Laterality of throwing by wild chimpanzees (n=16) of Mahale’s M group.

Name
AL
BB
CD
CE
CT
DE
DG
DW
FN
IV
MC
OR
OS
PM
PR
XM
TOTAL

Year of
Birth
1982
1981
1991
1998
1985
1963
1981
1988
1978
1993
1996
1991
1998
1988
1991
1995

Total
Throws
46
37
34
11
32
27
21
36
70
20
64
47
14
16
72
9
556

Both Hands

Left Hand

Right Hand

1
2
1
0
1
17
3
2
13
1
9
7
0
1
4
1
63

21
26
18
3
6
3
10
17
27
4
28
10
5
10
35
3
226 (46%)

24
9
15
8
25
7
8
17
30
15
27
30
9
5
33
5
267 (54%)

R vs. Lp<0.05
L

R

R
R

L = stat. sig. left bias; R = stat. sig. right bias; blank = not stat. sig. diff.
All chimpanzees are males, except for IV (bolded).

DISCUSSION

These results clearly contrast with those from themost thorough study done in captivity: Hopkins et al.7
found that of 89 subjects with six or more data-points, 50
were right-biased, 23 were left-biased, and 16 were unbiased. We have no explanation of this remarkable difference, although it joins a long list of behavioural contrasts
between nature and captivity, in which captive chimpanzees are lateralised, while wild chimpanzees are not(see summary in ref. 14). We can find no reports of twohanded throwing by captive chimpanzees; whether this isbecause such behaviour was not reported, or because it isabsent, remains to be seen.
Earlier, McGrew and Marchant15 hypothesised that
captive chimpanzees might be biased toward rightsidedness by their lengthy (sometimes lifetime) association with human caretakers, most of whom will have beenright-handed. However, a recent extensive study of four
great ape species (see below) found species-level rightsided laterality in captive chimpanzees, at least based on asingle bimanual measure16.
The implications for the evolutionary origins of human handedness are no clearer than the present state ofplay of contradictory studies of behavioural laterality.
Handedness in living Homo sapiens could be a derived
trait (of uncertain time-depth) or a primitive trait that
could date back as far as the last common ancestor of thepresent African apes and humans (perhaps as long ago as
6–7 million years). Results on manual laterality for the
only living Asian great ape, the orang-utan, Pongo pygmaeus, are confusing, as according to Hopkins et al.16,
they are a left-handed species.
What is clear is that chimpanzees both in captivity
and in nature habitually throw, despite being non-linguistic quadrupeds. Thus it is not necessary to invoke bipedal

stance or co-evolved language to explain the evolutionary
origins of throwing in the hominin lineage, and if the results from nature are to be believed, neither is it necessary
to link throwing to manual lateralisation.

ACKNOWLEDGEMENTS

We thank: the Tanzania Commission for Science andTechnology (COSTECH/UTAFITI), Serengeti (now Tanzania)
Wildlife Research Institute (SWRI/TAWIRI), Tanzania
National Parks (TANAPA) for authorisation to do the research; H.M. Nguli, E.T. Massawe, B.C. Mwasaga, and J.V.
Wakibara for special administrative assistance; M. Huffman,
K. Kawanaka, M. Nakamura, and S. Uehara for field collaboration; Philip and Elaina Hampton Fund (Miami University)
for financial assistance to LFM & WCM, and Japanese
MEXT #12375003, #16255007 for financial assistance to TN.
Toshisada Nishida died on 7 June, 2011 and is much missed.

REFERENCES

1. Calvin WH 1983. The Throwing Madonna: Essays on the
Brain. McGraw-Hill, New York.
2. Isaac B 1987. Throwing and human evolution. Afr Archaeol
Rev 5:3–17.
3. Goodall J 1964. Tool-using and aimed throwing in a community of free-living chimpanzees. Nature 201:1264–1266.
4. Köhler W 1927. The Mentality of Apes. 2nd edit. Kegan
Paul, Trench, Trubner, London.
5. Marchant LF 1983. Hand Preferences among Captive
Island Groups of Chimpanzees. PhD dissertation, Rutgers,
The State University of New Jersey, New Brunswick, NJ.
6. Hopkins WD, Bard KA, Jones A, Bales SA 1993. Chim­
panzee hand preference in throwing and infant cradling:
Implications for the origin of human handedness. Curr
Anthropol 34:786–790.
7. Hopkins WD, Russell JL, Cantalupo C, Freeman H,
Schapiro SJ 2005. Factors influencing the prevalence andhandedness for throwing in captive chimpanzees (Pantroglodytes). J Comp Psychol 119:363–370.
8. Hopkins WD, Russell JL, Schaeffer JA 2012. The neuraland cognitive correlates of aimed throwing in chimpanzees: A magnetic resonance image and behavioural study

Pan Africa News, 19(2), December 2012
on a unique form of social tool use. Phil Trans Roy Soc B
367:37–47.
9. Forrester GS, Quaresmini C, Leavens DA, Mareschal D,
Thomas MSC 2013. Human handedness: An inherited evolutionary trait. Behav Brain Res 237:200–206.
10. Nishida T (ed) 1990. The Chimpanzees of the Mahale
Mountains: Sexual and Life History Strategies. Universityof Tokyo Press, Tokyo.
11. Nishida T 2012. Chimpanzees of the Lakeshore. Natural
History and Culture at Mahale. Cambridge University
Press, Cambridge.
12.Nishida T, Zamma K, Matsusaka T, Inaba A, McGrew WC
2010. Chimpanzee Behavior in the Wild: An Audio-visual
Encyclopedia. Springer, Tokyo.
13. Martin P, Bateson P 2007. Measuring Behaviour: An
Introductory Guide. 3rd edit. Cambridge University Press,
Cambridge.
14. McGrew WC, Marchant LF 1997. On the other hand:Current issues in and meta-analysis of the behavioral laterality of hand function in nonhuman primates. Yearb Phys
Anthropol 40:201–232.
15. McGrew WC, Marchant LF 2001. Ethological studyof manual laterality in the chimpanzees of the Mahale
Mountains, Tanzania. Behaviour 138:329–358.
16. Hopkins WD, Phillips KA, Bania A, Calcutt SE, Gardner
M, Russell J, Schaeffer J, Lonsdorf EV, Ross SR, Schapiro
SJ 2011. Hand preferences for coordinated bimanual actions in 777 great apes: Implications for the evolution ofhandedness in Hominins. J Hum Evol 60:605–611.

<NOTE>

Playful Drumming by Immature
Wild Chimpanzees at Mahale: Do
They Enjoy Making Sounds?
Takahisa Matsusaka
Kansai University, Japan(E-mail: matsusaka.t@gmail.com)

INTRODUCTION

Before the Mahale Mountains National Park was established in 1985, several villages of Tongwe people lived
in the forest. Although many years have passed since thepeople left, we can still sometimes find the remains ofhuman artifacts on the forest floor while observing chimpanzees. Here, I report the rare cases where immature
chimpanzees found and drummed on clay pots discarded
by the former villagers and discuss their playful nature in
relation to the origin of music.

OBSERVATIONS(1) Case of an 11-Year-Old Adolescent Male, Cadmus (CD)
(25 Nov. 2002)
While CD was playing with a 4-year-old male infant(OS) in a thicket, CD found at 10:13:59 a clay cooking
pot (about 40-cm diameter, 30-cm high, with a 14-cm diameter hole at the bottom). He lifted it with his left handand approached OS with a play-face. OS approached andtouched CD, but soon returned to his mother to suckle.
At 10:15:29, an estrous adolescent female (QA)
showed leaf-clipping courtship near CD, who did not
respond. At 10:15:39, CD lifted the pot with both hands,
put it on his head like a helmet, and tapped 31 times on its

23
side with his right palm. Then he stamped on the groundfour times and pounded the pot three times with his left
fist. After stamping on the ground twice, he again tappedon the pot’s side with his right palm 14 times (Figure 1).

Figure 1. CD puts a pot on his head and taps it with hispalm.

At 10:16:42, CD lowered his head, placed the poton the ground, and kicked its rim twice with his rightheel. Then he lowered one side of the rim with his righthand and kicked the rim with his right heel ten times. At
10:17:19, CD stood up, kicked backward, and pushed the
pot with his right sole. The pot rolled off, and he left.
In total, CD drummed 60 times: 45 times with hisright palm, three times with his left fist, and 12 times withhis right heel. He calmly drummed with no pilo-erection.
His face was not visible when he put the pot on his head,but he drummed on it on the ground with no facial expression. He generally tapped lightly and did not make loudsounds. Although he might have heard the pot’s large resonance from inside it, the sounds did not seem to attract
others. No one approached CD during his drumming,
even though six chimpanzees rested within 10 m: OS andhis mother, a 1-year-old male infant (TD) and his mother,an adult male (DG), and an adolescent female (QA).
(2) Case of a 6-Year-Old Juvenile Male, Michio (MC) (14Feb. 2003)
At 10:46:56, while MC was walking in a thicket, hefound a large clay water pot that was about 50-cm high,with a 45-cm diameter, a narrow neck, and an 18-cm
diameter mouth. He slapped the bottom of it three times
with his left palm, rolled it, and slapped it once with hisright palm. Then he stepped onto it with both feet, soon
got off it, lowered the pot’s mouth to the ground, andslapped its bottom with his left palm. He then pushed
and rolled it, slapped it simultaneously with both palms,
and slapped it nine times with both hands in turns. At
10:47:19, he stopped but soon started playing again invarious ways (Figure 2, Video 1: available online atmahale.main.jp/PAN/19_2/19(2)_05.html): slapping the
pot with his palms, pushing and rolling it, stamping on itwith his soles, kicking it with his heels, stepping up onto

24

Pan Africa News, 19(2), December 2012
male infant, TD and his mother, and two adult males (AL,PM). Only TD came to the spot after the pot was finallysmashed.

DISCUSSION

Figure 2. MC slaps a pot with his palms.

it, dropping his buttock on it, and lying supine on it. He
also showed such exploratory behaviors as touching its
crack, looking into it, putting his arm into it, etc.
At 10:52:11, MC strongly kicked the pot five times
with his right heel, chipping off a small piece. He touched
the spot with his right hand and slapped it eight times
with both palms in turns. Then he stepped up onto the
pot and stamped on it six times with his sole, and the pot
finally broke into pieces, and he fell off of it (10:52:57).
One-year-old male infant, TD, approached within 5 m of
MC and observed the scene. MC lifted one of the broken
pieces, but soon went away and approached TD and his
mother. After a while, TD came, touched the broken pieces, and stepped on them, but no one paid any attention to
the pieces after that.
In total, MC drummed on the pot 199 times: 36 times
with his right hand, 112 times with his left hand, fourtimes simultaneously with both hands, 25 times with hisright leg, eight times with his left leg, four times withhis buttock, and ten times with unidentified body parts.
He used his left hand more than his right hand and his
right leg more than his left leg. When he drummed withhis hands, he usually used his palms (149 times) but healso used his fist (twice) and his wrist (once). When hedrummed with his legs, he used his sole (23 times) and hisheel (10 times).
MC usually drummed on the side or the bottom of thepot, but he also hit its rim (once) and neck (twice). Since
the pot has a round bottom, it was unstable and easy toroll. MC seemed to try to control the pot’s orientation andposition by pushing or dragging it, lowering or liftingits mouth, and supporting it with his limbs. Supporting
behaviors were observed nine times: three times with hisright hand, once each with both hands, the right leg, theleft leg, both legs, and combinations of right hand/left
leg and right hand/right leg. His right hand was used six
times, and his left hand was used only once.
MC drummed the pot with no pilo-erection andwith a half-open mouth, which is his relaxed face. He
drummed relatively strongly, and the sounds were probably audible to others nearby. However, most chimpanzees
did not show much interest in his drumming; six others
rested within 10 m: MC’s mother and her 1-year-old fe-

CD put the pot on his head like a helmet and tapped itwith his palms and kicked it on the ground. He drummed
60 times in about two minutes. MC showed more varied
behavioral patterns, including slapping, kicking, stepping up, and rolling. He drummed 199 times in about six
minutes. These can be regarded as “object play” in which
they experienced various types of object manipulation,
and also as “sound play” in which they enjoyed making
sounds. During drumming, they sometimes changed the
pot’s orientation and position and drummed on various
spots with different body parts. They seemed to enjoy
modifying the sounds and hearing the unexpectedly large
and resonant sounds from the pot.
Immature chimpanzees sometimes make sounds
during play. They might slap a tree buttress or walk on
branches in an exaggerated style to make pitapat sounds.
They also make sounds when they play with dry leaves1.
Some immature chimpanzees (including CD and MC) occasionally slap their own bellies with their palms to makespanking sounds as solo play or play solicitation 2,3. They
may also enjoy making sounds in these cases, but thesecases generally last only for a few seconds. The cases
observed in this study were exceptionally long. The relatively large and resonant sounds of the pot should haveencouraged them to continue drumming.
Adult male chimpanzees often hit or kick a substratum to make banging sounds as an intimidation display,such as slapping the ground or slapping/kicking tree buttresses or metal walls4,5. The clay pot drumming observedin this study featured several differences from such adultdrumming. CD and MC were not very excited, unlike
adult displays; they did not show pilo-erection or suchtense facial expressions as compressed lips. They seemed
relaxed and drummed for a much longer duration. They
did not pay much attention to other chimpanzees and appeared absorbed in their solo activity. Others did not showmuch interest in their drumming, either.
An individual difference was found in the laterality.
CD usually drummed with his right hand and leg, while
MC used his left hand and right leg more often. Whensupporting the pot, MC mostly used his right hand. Thus,
MC showed a tendency of role differentiation in using
his hands for drumming and supporting. Individual differences of drumming patterns and laterality were also
reported for intimidation drumming by adult males4,5.
In summary, two immature chimpanzees engaged inlong and variable solo play, suggesting that they enjoyedproducing sounds. In discussion on the origin and evolution of music, much attention has focused on such social
aspects as vocal communication and sexual selection6.
However, the individual capacity to enjoy manipulating
sounds also seems essential for the origin of music. This
study suggests that chimpanzees have the potential toenjoy making sounds and supports the notion that manual
drumming in African great apes is homologous to human
instrumental music7.

Pan Africa News, 19(2), December 2012
ACKNOWLEDGEMENTS

I thank the Tanzania Commission for Science andTechnology, the Tanzania Wildlife Research Institute, and the
Tanzania National Parks for permission to work at Mahale,
the Mahale Mountains Wildlife Research Centre and Mahale
Mountains National Park for logistic support, and Dr. H. Hayaki
and Dr. M. Nakamura for their valuable comments on this manuscript. This study was financially supported by MEXT Grant
in-Aid for Scientific Research (#12375003 and #16255007 to T.
Nishida)REFERENCES

1. Nishida T, Wallauer W 2003. Leaf-pile pulling: An unusual play pattern in wild chimpanzees. Am J Primatol
60:167–173.
2. Nishida T, Matsusaka T, McGrew WC 2009. Emergence,
propagation or disappearance of novel behavioral patterns in the habituated chimpanzees of Mahale: A review.
Primates 50:23–36.
3. Nishida T, Zamma K, Matsusaka T, Inaba A, McGrew WC
2010. Chimpanzee Behavior in the Wild: An Audio-Visual
Encyclopedia. Springer, Tokyo.
4. Arcadi AC, Robert D, Boesch C 1998. Buttress drummingby wild chimpanzees: Temporal patterning, phrase integration into loud calls, and preliminary evidence for individual distinctiveness. Primates 39:505–518.
5. Nishida T 2003. Individuality and flexibility of cultural
behavior patterns in chimpanzees. In: Animal Social
Complexity. de Waal FBM, Tyack PL (eds), Harvard
University Press, Cambridge, MA, pp. 392–413.
6. Wallin NL, Merker B, Brown S (eds) 2000. The Origins ofMusic. A Bradford Book, Cambridge, MA.
7. Fitch WT 2006. The biology and evolution of music: A
comparative perspective. Cognition 100:173–215.

<NOTE>

Comparison of the Longevity of
Chimpanzee Beds between Two
Areas in the Mahale Mountains
National Park, Tanzania

25
METHODSThis study was conducted in Mahale Mountains
National Park, Tanzania. This site has a rainy season fromearly October to mid-May7. We focused our research in
two areas in the park, Kasoje and Miyako (Figure 1). The
Kasoje area is composed of gallery forest 8,9, and is thecenter of the range of the M group of chimpanzees. The
Miyako area consists of Brachystegia woodland withgallery forest along riverside area8,9, and is occupied bychimpanzees of the Y group10.
Across both sites, we found 102 fresh beds from 3
October 2006 to 16 February 2007, and classified them by
their date of construction (Table 1). Of these, we selected
70 beds to estimate bed longevity, and 32 beds to study
bed construction. For the longevity study, we classified
beds into two categories based on their time of construction. Beds constructed during Period 1 (3 to 10 October)were monitored until 24 May 2007. Beds constructedduring Period 2 (11 October to 10 November) were monitored for a minimum of 100 days. Beds constructed after
Period 2 were not included in analyses.
Beds were monitored approximately once per week(mean interval = 8.7 days, range = 6–28 days). KZ monitored the beds from 3 October 2006 to 16 February 2007,
and MM monitored them from 17 February to 24 May
2007. We defined bed leaves as decayed when all of leaves
had fallen from the branches in the bed 3,5, and definedbeds as decayed when they were no longer clearly discernable as chimpanzee beds1,2,4–6.
We monitored 9 beds in the Kasoje area and 7 bedsin the Miyako area during Period 1. During Period 2, wemonitored 38 beds in Kasoje area. The minimum distance
between bed sites of the two areas was about 2 km (Figure 1).

Koichiro Zamma1,2 & Masayuke
Makelele2
1 Great Ape Research Institute, Hayashibara, Japan
2 Mahale Mountains Chimpanzee Research Project(E-mail: zamma@gari.jp)

INTRODUCTION

Chimpanzee (Pan troglodytes) beds have been used
as an indicator for the presence of chimpanzees, and thelongevity of beds is one of the important variables for
estimating chimpanzee population sizes1,2. The longevityof chimpanzee beds can differ due to location, vegetation,and season3–6. For example, beds in forested sites decay
faster than those in woodlands5, and beds in the dry season decay faster than those in the rainy season3,5.
A previous study 6 on the longevity of chimpanzee
beds was conducted in the Mahale Mountains National
Park, Tanzania. Because the study6 was conducted only inthe forested area in the dry season, we studied chimpanzee beds both in forested and woodland sites during therainy season.

Figure 1. Locations of chim­panzee beds in Kasoje (closed
circle) and Miyako (open cir­cle) during period 1.

Pan Africa News, 19(2), December 201226
Table 1. Tree species used for chimpanzee bed.
Area

Local name
(Kitongwe)

Kasoje Lulumasha*
Mkibu gwesimbwa*
Buhono*MjohoroIbonobono*
Msabasaba
Mnyoni
Kafulujege*
Kahwibili
Kamilaninga*
Mjohoro (dead)§
Mlama*
Mlyansekesi*
Bulonje*Ikubila*Mtelele*MuhokoBuhono* & Kahwibili†
Buhono* & Lulumasha*†
Buhono* & Mlama*†
Buhono* & Bulonje*†
Kahwibili & Mnyoni†
Kahwibili & Lulumasha*†
Kahwibili & Mgongogongo*†
Miyako Mlonje*
Kashindabilangulube
Lisalike
Mkusi
Total

Scientific name

Family name

Number of beds
Total

Pycnanthus angolensis #1
Cordia millenii
Pseudospondias microcarpa #2
Senna spectabilis #3
Croton sylvaticus
Syzygium sp.
Bridelia micrantha #4
Lecaniodiscus fraxinifolius
Xylopia parviflora #5
Bridelia atroviridis#3
Combretum molle #6
Synsepalum brevipes
Dracaena usambarensis #7
Ficus sur
Stereospermum kunthianum
Aporrihiza paniculata#2 & #5#2 & #1#2 & #6#2 & #7#5 &#4#5 & #1#5 & Anthocleista schweinfurthii
Mimusops bagshawei
Blighia unijugata?
Croton megalocarpus

Myristicaceae
Boraginaceae
Anacardiaceae
Fabaceae
Euphorbiaceae
Myrtaceae
Euphorbiaceae
Sapindaceae
Annonaceae
Euphorbiaceae
Combretaceae
Sapotaceae
Dracaenaceae
Moraceae
Bignoniaceae
Sapindaceae

Loganiaceae
Sapotaceae
Sapindaceae?
Euphorbiaceae

19
18
13
6
5
5
3
3
3
2
2
2
2
1
1
1
1
2
1
1
1
1
1
1
4
1
1
1
102

After
Period 1 Period 2 Period
2
7

1

10
12
6

5
3

3
6
1

2

2
1

1

1
1

1

4
1
1
1
16

38

16

*: Fruits, leaves, and/or piths of this plant were eaten by chimpanzees.
†: Two species of tree were used to make a bed.
§: Two beds were constructed at the base of dead trees but were built from living vines including Saba comorensis (Apocynaceae), Keetia
venosa (Rubiaceae), and Tetracera potatoria (Dilleniaceae).

RESULTSWe recorded 21 tree species that were used for constructing chimpanzee beds (Table 1). The most frequently
used species was Pycnanthus angolensis, followed by
Cordia millenii and Pseudospondias microcarpa. Manyof the trees in which beds were constructed (78/102) are
known food trees of Mahale chimpanzees. Eight beds
were also found in the invasive tree, Senna spectabilis,
and two of them were constructed at the base of deadtrees.
The mean longevity of chimpanzee beds during
Period 1 was 87 days (n = 9, range = 21–231) in the
Kasoje area and 164 days (n = 7, range = 13 to > 233) inthe Miyako area (Figure 2). Three beds in the Miyako
area were not decayed by the end of the survey period.
Bed longevity was significantly longer in Miyako than in
Kasoje (Mann-Whitney U test, n1 = 9, n2 = 7, U = 12, p <0.05).
During Period 2, we monitored 38 chimpanzee bedsin Kasoje. We set a 100 day limit for monitoring thesebeds and more than half of the beds (22/38) were decayedby the end of this period (Figure 3). This was not statistically different from the decay rate for Period 1 beds in
Kasoje over 100 days (6/9; Fisher’s exact test p = 0.72),
but was significantly greater than the decay rate for bedsin Miyako (1/7; Fisher’s exact test, p < 0.05).
The mean time for leaf decomposition in beds was
76 days in Kasoje (n = 9, range = 14–158) and 126 days in
Miyako (n = 7, range = 13–157). These numbers were notstatistically different according to a Mann-Whitney U test
(Figure 4, n1 = 9, n2 = 7, U = 15, p = 0.088).

DISCUSSION

There are two methods for measuring the longevityof chimpanzee beds: the time until leaves decays and thetime until the bed structure decays1–6. Bed leaves decayed
faster in Kasoje in the dry season (49 days, n = 41, range= 18–110)6 than in the rainy season (76 days) in this study.
In tree species of bed no remarkable differences could beseen between this and previous study, and Pycnanthus
angolensis was the most frequently used species in bothstudies. Leaves in the dry season are thought to havePan Africa News, 19(2), December 2012

Figure 2. Rate of decay of chimpanzee beds in period 1.
*: Three beds were remained undecayed.

27

Figure 3. Rate of decay of chimpanzee
beds in Kasoje in Period 2.

Figure 4. Rate of decay of bed leaves in Period 1.

lower moisture content and, thus, decay faster3. The resultin this study is consistent with previous studies3,5.
In contrast, the longevity of bed structures in Kasoje
was longer in the dry season (131 days, n = 41, range =30–189)6 than in the rainy season (87 days) in this study.
This indicates that different mechanisms are relevant
for the decay of leaves in beds than for decay of the bed
structure. Over the period of this study (3 October 2006
–24 May 2007) 1,562 mm of precipitation fell and almost
two-thirds of the total precipitation (1,001 mm) fell during the first 3 months (Mahale Mountains Chimpanzee
Research Project data). We propose a hypothesis: because
dead leaves often remain in branch structure in beds andabsorb rainfall during the rainy season, the increased
weight of the wet branches may contribute to the deterioration of the bed structure.
The longevity of chimpanzee beds in Miyako was
longer than that in Kasoje, but this difference cannot be explained by rainfall because the amount of precipitation did
not differ between the two areas11. The longevity of chimpanzee beds in woodlands in the Ugalla area, Tanzania islong (more than 358 days (Yoshikawa M, personal communication); 432 days5), and beds in Miyako area also

constructed in woodland. The strength of branches of tree
species in woodland may be a key factor contributing to
the longevity of chimpanzee beds in these areas.
The present study and previous studies (e.g. ref. 1–6)
indicate that the longevity of chimpanzee beds differs
according to season and vegetation type. These studies
indicate the need for broader range of bed longevity or for
appropriate estimates of bed longevity in each site to accurately estimate chimpanzee population sizes.

Acknowledgements

We thank TAWIRI, TANAPA and COSTECH for permission to do the field research, and MMNP and MMWRC for logistic support. We also thank T. Sakamaki for providing information about the beds of Y group chimpanzees, H. Bunengwa,
M. Mwami, H. Ihobe, N. Itoh, H. Ogawa, and M. Yoshikawa for
their help and advices, the members of MMCRP for providing
precipitation data, and M. Kiyono and C. Tokimatsu for data
handling. This study was financially supported by the Global
Environment Research Fund of the Ministry of Environment,
Japan (F-061 to T. Nishida).

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<BOOK INFO>

Wild Cultures: A Comparison
between Chimpanzee and Human
Cultures
By Christophe Boesch
How do chimpanzees say, “I want to have sex withyou?” By clipping a leaf or knocking on a tree trunk?
How do they eat live aggressive ants? By using a shortstick with one hand or long stick with both? Ivorian andTanzanian chimpanzees answer these questions differently, as would humans from France and China if asked
how they eat rice. Christophe Boesch takes readers into
the lives of chimpanzees from different African regions,highlighting the debate about culture. His ethnography
reveals how simple techniques have evolved into complex
ones, how teaching styles differ, how material culture
widens access to new food sources and how youngsters
learn culture. This journey reveals many parallels between humans and chimpanzees and points to striking
differences. Written in a vivid and accessible style, Wild
Cultures places the reader in social and ecological contexts that shed light on our twin cultures.

Pan Africa News, 19(2), December 2012296 pages
Cambridge University Press
Publication date: September 2012
Hardcover $95.00
ISBN: 9781107025370
Contents
Introduction
1. Studying culture in the wild
2. From human culture to wild culture
3. Shaping nature into home: about material culture
4. One for all and all for one: about social culture
5. I want to have sex with you: about symbolic culture
6. Learning culture: from pupils to teachers
7. Dead or alive? Towards a notion of death and empathy
8. Wild culture – wild intelligence
9. Uniquely chimpanzee – uniquely human
Epilogue: will we have the time to study chimpanzee
culture?
References
Index

Pan Africa News, Vol. 19, No.2
Published in December, 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)