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 2013
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, Kyoto University, Japan
Agumi Inaba, Mahale Mts. Chimpanzee Research Project
Eiji Inoue, Kyoto University, Japan

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VOL. 20, NO. 2

Contents<ARTICLE>
Chimpanzee (Pan troglodytes schweinfurthii)
Population Density and Abundance inKibira National Park, Burundi
Dismas Hakizimana & Marie-Claude Huynen

16<NOTE>
Fission-Fusion in Chimpanzees: Feeding asa Proximal Mechanism at Gombe
Andrea Blackburn & William C. McGrew

19<NOTE>A Juvenile Chimpanzee Played with a Live
Moth
Michio Nakamura			

22<NEWS>
The 1st MWCS Scholarship Student Awarded
DiplomaKazuhiko Hosaka
With a report from Butati R. Nyundo “My teachingexperience at my hometown”			
24<BOOK INFO>
Primates and Cetaceans: Field Researchand Conservation of Complex MammalianSocieties
Edited by Juichi Yamagiwa and Leszek Karczmarski

25

Deadline of the next issue is April 2014!

15

Pan Africa News, 20(2), December 201316

<ARTICLE>

Chimpanzee (Pan troglodytes
schweinfurthii) Population
Density and Abundance in Kibira
National Park, Burundi
Dismas Hakizimana1,2 & MarieClaude Huynen1
1 Department of Behavioral Ecology, University of Liege,
Belgium
2 Department of Biology, University of Burundi, Burundi(E-mail: hakdismas@yahoo.fr)

INTRODUCTION

The eastern chimpanzee, Pan troglodytes schweinfurthii, is an “endangered” chimpanzee subspecies ofglobal conservation concern, living at low density (lessthan 1 individual/km²) in most of its range (Keele et al.
2006; Plumptre et al. 2011) in eastern Africa. As chimpanzees have a relatively low reproductive rate (Caldecott& Miles 2009), their ability to recover from population
decline is limited. Documenting the extent and severity ofsuch decline events, as well as targeting adequate conservation plans, requires reliable baseline data on density estimates of great apes (Oates 1996) in area of occurrence.

In Burundi, eastern chimpanzees are mostly reported
to occur from the North-Western Kibira National Park
(KNP), which is contiguous with Nyungwe National Park
(NNP) in Rwanda, both forming one continuous landscape (Figure 1). Unfortunately, during the 1993–2007
conf lict, KNP was permanently occupied by armed
groups and its biodiversity has been extensively destroyed
by armed gangs’ progression, bush fires, poaching, illegal
logging and clearing for agriculture. The purposes of oursurvey were to provide reliable information on chimpanzee abundance in the KNP and to identify sectors whereconservation efforts should be prioritized.

METHODS

Study area
KNP is located in the North-West of Burundi on theCongo-Nile Divide, between 2°36'52'' and 3°17'08" south
latitude and between 29°13'31'' and 29°39'09" east longitude. It covers 400 km² of montane forest, lying from1,600 to 2,666 m of altitude (Arbonier 1996). Relief, moremarked on the western side, is characterized by steep
slopes on both sides of the Congo Nile Divide. The park
is shaped into four sectors, from south to north: Teza,
Musigati, Rwegura and Mabayi, covering respectively
57.94 km², 154.24 km², 124.23 km² and 63.59 km² (Figure
2).

Figure 1. Location of Kibira and Nyungwe National parks.
(Reproduced from Africa: Atlas of Our Changing Environment by UNEP (2008).)

Pan Africa News, 20(2), December 2013

17

Figure 2. The four sectors of the KNP with location of the research transects.

Data collection
Between September 2011 and February 2013, we repeatedly surveyed sixteen linear transects of 3 km each,to census chimpanzee populations density and abundance
using the marked nest count method (Kouakou et al.
2009). The inter-visit interval was of three weeks, andeach transect was censused eighteen times over the entire
study period.
We walked along the census transects with two fieldassistants at an average speed of 0.7 km per hour to ensure
detection of all chimpanzee nests. During the first walk,
we marked all existing nests with a piece of flagging tapeattached to nesting trees (Plumptre & Reynolds 1996).
During the following visits, only new nests built since
the last passage were recorded and marked. For each nest

we systematically recorded the perpendicular distance totransect line, GPS position and altitude.
Data analysis
To estimate chimpanzee density for the whole park,
we combined nest counts from all transects. We used the
DISTANCE software (version 6.0) (Thomas et al. 2009)
to obtain chimpanzee density estimates. According to
Buckland et al. (2001), 5% of observations occurring at
the furthest distance from transect were truncated. Fourmathematical key (Uniform, Half-normal, Hazard rateand Negative exponential) modeling nest detection function were tested, in combination with 3 adjustments terms(Cosine, Simple polynomial and Hermite polynomial).
We calculated Akaike’s Information Criterion (AIC) forPan Africa News, 20(2), December 201318
each model and the best model had a lowest AIC value(Buckland et al. 2001). Nest-building chimpanzee density
was estimated by following Plumptre & Reynolds (1996).

RESULTSA total of 864 km of line transects were surveyed and
we recorded 471 new nests of chimpanzees, at an altitude
ranging between 1,956 – 2,552 m.
Half-normal key with simple polynomial adjustment
was the model fitting the best our distribution of perpendicular distances. Chimpanzee density estimate in theentire park was 0.509 individual/km² (95% CL: 0.305 –0.847, CV: 24.37%), which corresponds to a total population size of 204 weaned individuals (95% CL: 122 – 339).
Chimpanzee densities for each sector of the park are given
in Table 1.
Mabayi has an important chimpanzee density compared with other sectors. Its chimpanzee density is fourtimes greater than Rwegura, six times greater thanMusigati and three times greater than Teza.

DISCUSSION

Chimpanzee density and abundance in the Kibira NationalPark
Our results are markedly lower than previous estimates by Barakabuye et al. (2007) who reported a density
estimate of 0.984 individual/km² (95% CL: 0.601 – 1.612),
and a total population of 394 chimpanzees (95% CL: 240– 645). This discrepancy is likely due to different sampling efforts. First, our estimated chimpanzee density isbased on 18 months of survey while Barakabuye et al.
(2007) survey only covered a three-month period. Shorttime survey provides less reliable information, especiallyfor P. troglodytes known to use large home ranges (Johns& Skorupa 1987). In order to take into account seasonalvariations in behavior, Baldwin et al. (1982) suggested
a minimum of one year for such survey. Second, we applied an equal sampling effort throughout the four sectorsof the KNP while Barakabuye et al. (2007) placed halfof their transect lines in the sole Mabayi sector, which
is most likely the one harboring the largest chimpanzee
population. Extrapolation of their estimates to the entire
park logically leads to an overestimation (Buckland etal. 2010). The longer time we devoted to data collection
and the use of the marked nest count method leads us tosay that our estimate would be more representative of the
KNP.

The lower estimate of the chimpanzee population sizein our study is in logical agreement with the region recent
history. Biodiversity of the KNP suffered from several
years of armed conflict. Armed groups practiced poaching and subsistence hunting. As chimpanzee is the largest
animal of the park, it is evident that it was a war victim.
Comparison within sectors
Determining the variation of distribution of chimpanzees between given areas is very important for researchers and conservationists (Morgan et al. 2006). Our results
show that chimpanzees are more abundant in the Mabayi
sector, directly contiguous with the NNP in Rwanda. Thismight be because this area was less affected by armed
conflicts compared to Teza and Musigati which were permanently occupied by rebels. Another reason might bethat some tree species primordial as food sources and nest
supports for chimpanzees, are only or more represented inthis part of the park (Hakizimana, unpublished data).
Comparison within Kibira and Nyungwe National Parks
Kibira and Nyungwe are two contiguous nationalparks, but Nyungwe (1,080 km²) is twice larger than
Kibira (400 km²). Barakabuye et al. (2007) found achimpanzee density much higher in the Kibira compared to Nyungwe (0.984 individuals/km², 394 individuals in Kibira; 0.353 individuals/km², 382 individuals in
Nyungwe). This is a puzzling difference, as Nyungwe isknown to have been better protected (Barakabuye et al.
2007). If confirmed, the highest chimpanzee density theyfound in Kibira might result from the high density of tree
species on which chimpanzees rely heavily (Balcomb et
al. 2000; Tweheyo et al. 2004). To ensure a robust comparison and solve ambiguities, a rigorous study should be
led by the same team in both parks focusing on census
and resource availability.
Conservation implications
Intensive effort must be devoted to the chimpanzee’s
long-term conservation throughout their range. Researchon the species is still young in Burundi compared toneighboring countries. Therefore, it is very important
to implement a monitoring program of chimpanzee
populations to constitute a database allowing detecting
population trends. Given the low rate reproduction ofchimpanzees, Kühl et al. (2009) suggested that an inventory should be repeated every one to five years in order toTable 1. Population density and abundance, with confidence limits (CL) of nest-building chimpanzees in the four
sectors composing the Kibira National Park.

Location

Area (km²)

Estimated
density

95% CL

Number ofindividuals

95% CL

Teza

57.94

0.352

0.159 – 0.778

20

9 – 45Musigati

154.240.190

0.093 – 0.390

29

14 – 60

Rwegura124.230.293

0.106 – 0.812

36

13 – 101

Mabayi

63.59

1.172

0.744 – 1.847

75

47 – 117

Entire park

400

0.509

0.305 – 0.847

204

122 – 339

Pan Africa News, 20(2), December 2013
quickly detect population changes. Also, further studies
must emphasize more detailed data on tree phenology thatchimpanzees rely on. As Nyungwe and Kibira National
Parks are contiguous, an effective transboundary strategicplan to conserve both parks as one landscape is necessary
to maintain viable population of chimpanzees.

ACKNOWLEDGMENTS

Financial support for this work came from Burundian
Gover nment, with additional funds being provided byPatrimoine de l’Université de Liège, International Foundationfor Science (IFS) and Ecole régionale post-universitaired’aménagement et de gestion intégrée des forêts tropicales(ERAIFT) via WBI program. Thanks are due to the guards of
KNP, without whom this work could not have been carried out.
We thank Nicolas Granier, Charles-Albert Petre, and PhilippeTamini for their helpful cooperation. We are grateful to Dr.
Sandra Tranquilli for her helpful comments on the manuscript.

REFERENCES

Arbonier M 1996. Parc National de la KIBIRA: Plan deGestion. INECN/CIRAD-FORET.
Balcomb SR , Chapma n CA , Wr a ng ha m RW 20 0 0.
Relationship between chimpanzee (Pan troglodytes) density and large, fleshy-fruit tree density: conservation implications. Am J Primatol 51:197–203.
Baldwin PJ, McGrew WC, Tutin CEG 1982. Wide-ranging
chimpanzees at Mt. Assirik, Senegal. Int J Primatol
3:367–385.
Barakabuye N, Mulindahabi F, Plumptre AJ, Kaplin
K, Munanura I, Ndagijimana D, Ndayiziga O 2007.
Conservation of Chimpanzees in the Congo Nile Divide
forests of Rwanda and Burundi: Unpublished Report. No
98210-G-GO95/GA 0282. Arlington VA: US Fish andWildlife Service (USFWS).
Buckland ST, Anderson DR, Burnham KP, Laake JL,
Borchers DL, Thomas L 2001. Introduction to DistanceS a m pli ng: E st i m a t i ng Ab u n d a n c e of Biolog i c al
Populations. Oxford University Press, Oxford.
Buckland ST, Plumptre AJ, Thomas L, Rexstad EA 2010.
Design and analysis of line transect surveys for primates.
Int J Primatol 31:833–847.
Caldecott J, Miles L 2009. Atlas Mondial des Grands Singes
et de leur Conservation. UNESCO, Paris.
Johns AD, Skorupa JE 1987. Responses of rain-forest
primates to habitat disturbance: review. Int J Primatol
8:157–191.
Keele BF, Heuverswyn FV, Li Y, Bailes E, Takehisa J,
Santiago ML, Bibollet-Ruche F, Chen Y, Wain LV,
Liegeois F, Loul S, Ngole EM, Bienvenue Y, Delaporte E,
Brookfield JFY, Sharp PM, Shaw GM, Peeters M , Hahn
BH 2006. Chimpanzee reservoirs of pandemic and nonpandemic HIV-1. Science 313:523-526.
Kouakou CY, Boesch C, Kuehl H 2009. Estimating chimpanzee population size with nest counts: validating methods inTaï National Park. Am J Primatol 71:447–457.
Kühl H, Maisels F, Ancrenaz M, Williamson EA 2009.
Lignes Directrices pour de Meilleures Pratiques en
Matière d’Inventaire et de suivi des Populations de Grands
Singes. Gland, Suisse : Groupe de spécialistes des primatesde la CSE de l’UICN. 32 pp.
Morgan D, Sanz C, Onononga J R, Strindberg S 2006. Apeabundance and habitat use in the Goualougo Triangle,
Republic of Congo. Int J Primatol 27:147–179.
Oates JE 1996. African Primates: Status Survey and19Conservation Action Plan, Revised Edition. IUCN/SSC.
Plumptre AJ, Reynolds V 1996. Censusing chimpanzees in
the Budongo forest, Uganda. Int J Primatol 17:85–99.
Plumptre AJ, Rose R, Nangendo G, Williamson EA, Didier
K, Hart J, Mulindahabi F, Hicks C, Griffin B, Ogawa H,
Nixon S, Pintea L, Vosper A, McClennan M, Amsini F,
McNeilage A, Makana JR, Kanamori M, Hernandez A,
Piel A, Stewart F, Moore J, Zamma K, Nakamura M,
Kamenya S, Idani G, Sakamaki T, Yoshikawa M, Greer D,
Tranquilli S, Beyers R, Furuichi T, Hashimoto C, Bennett
E 2011. Chimpanzé de Schweinfurth (Pan troglodytes schweinfurthii) : État de Conservation de l’Espèce et Pland’Action 2010–2020. Groupe de spécialistes des primatesde la CSE/UICN, Gland, Suisse.
Thomas L, Laake JL, Rexstad E, Strindberg S, Marques
FFC, Buckland ST, Borchers DL, Anderson DR, Burnham
KP, Burt ML, Hedley SL, Pollard JH, Bishop JRB,
Marques TA 2009. Distance 6.0. Release 2. Research Unit
for Wildlife Population Assessment, University of St.
Andrews, UK. http://www.ruwpa.st-and.ac.uk/distance/
Tweheyo M, Lye KA, Weladji RB 2004. Chimpanzee dietand habitat selection in the Budongo Forest Reserve,
Uganda. Forest Ecol Manag 188:267–278.

<NOTE>

Fission-Fusion in Chimpanzees:
Feeding as a Proximal
Mechanism at Gombe
Andrea Blackburn & William C.
McGrew
Department of Archaeology & Anthropology, University ofCambridge, U.K.
(E-mail: wcm21@cam.ac.uk)

INTRODUCTION

Fission-fusion is the species-typical and universal
social organisation of Pan troglodytes, as recognised
45 years ago by the pioneering field research of such researchers as Goodall (1968), Nishida (1968) and Sugiyama(1968). In fission-fusion, the group (or community), subdivides into temporary parties, which may further fragment or reunite, such that over the course of a day, anindividual may be solitary or sociable to varying degrees(Aureli et al. 2008). The standard variable for measuring
fission-fusion is party size, that is, the number of individuals in spatial association in any one period or point
in time. Explanations for variation in party size are many,
but the most prevalent one is feeding competition, so thatthe larger the party, the greater the competition. Thus,
individuals seeking to reduce or avoid competition willfission, either to forage alone or with fewer companions.
Following this line of argument, we hypothesise that party
sizes will be lower during bouts of feeding than before orafter feeding.

BACKGROUND

Wrangham (1977) was the first to posit a positive correlation between group size and feeding competition inPan Africa News, 20(2), December 201320
chimpanzees. (In the broad sense, ‘group’ indicates any
aggregation, in any species, primate or otherwise.) The
logic is simple: If resources are finite at any point in space
and time, then the more organisms seeking to harvest
those resources at the same time and place, the greater
will be the competition, all other things being equal. For
chimpanzees, the group size problem is ‘subcontracted’ toephemeral parties, yielding maximum flexibility in dayto-day social relations. At one extreme is solitary foraging, when resources are sparse, versus maximally social
foraging, in which party size is 100% of group size, whenresources are abundant. Many other variables also influence group size, such as predation, reproduction, aggression, rank, seasonality, demography, etc. (see Aureli et al.
2008, for the most comprehensive and ambitious attempt
to model fission-fusion).
For chimpanzees, many studies have found correlations between party size and proxy measures of feeding
competition (e.g. Matsumoto-Oda et al. 1998; NewtonFisher et al. 2000; Itoh & Nishida 2007). The usual study
design is to relate party size (however defined) to one ormore measures of food supply, such as fruit availability,
in terms of abundance or distribution of resources. These
measures of food supply are then related to various othermeasures, such as number of oestrous females, wet versus
dry season, etc. Most (but not all, see Hashimoto et al.
2003) studies have found the expected relationship: When
resources are scarce, chimpanzees range in smaller parties than when resources are abundant.

However, gaps exist in these studies. Party size (however defined) is a static (‘snapshot’) measure, sampled atvarious points in time over the course of a day. We havefound no studies based on focal sampling that describe thedynamics of the process, for example, tell us how manyfission-fusion transitions a chimpanzee makes in a day. Or
what factors (activity, age, sex, rank, etc.) prompt a chimpanzee to leave or join a party. Furthermore, although the
hypothesised causal variable is feeding competition, we
can find no published data that directly address this, either by contest or scramble. Most studies do not measure
feeding but instead rely on food, usually in terms of availability (versus, e.g., quality). Many studies of food supply
do not relate it to actual chimpanzee foraging, much lessto competition (e.g. Isabirye-Basuta, 1988). Thus, descriptive studies at this macro-level are indirect and correlative only (see Kummer 2008, Yamagiwa 2008, for similarcomments on the lack of ethological data).
We sought to tackle some of these issues at the sequential, dynamic, micro-analytical level, by comparing
party sizes when feeding versus non-feeding, in real-time
sequences, focussing on individual behaviour. Other studies have shown static differences in group size in relationto activity (e.g. Newton-Fisher, 1999), but none seems tohave made this basic comparison.) We chose to test the
hypothesis in the most sociable age-sex class of chimpanzees, independently-ranging males, because it is the mostchallenging one.

Table 1. Number of companions of independently-ranging male chimpanzees at Gombe, before, during and after boutsof feeding. See text for explanation of four stages.

30 min beforenmedianNumber of companions
Start feeding
Stop feedingnmediannmedian30 min afternmedian

EV

9

3

84

2

84

2

7

6

FB

15

7

68

4.5

68

5.5

17

7

FG

4

2

48

2.5

48

3

3

2

GB

5

2

62

2

62

2

6

2.5

HG

5

2

44

2

44

3

4

7

HM

7

4

53

2

53

2

7

3

JJ

7

4

57

2

57

2

5

7

MK

8

7

46

1

46

1.5

7

4

SH

4

6.5

46

0

46

0

7

7

ST

5

2

50

1

50

1

6

1.5

Total

69

-

558

-

558

-

69

-

Median

6

-

51.5

-

51.5

-

6.5

-

Subject

Median ofmedians

3.5

2

2

5

Pan Africa News, 20(2), December 201321

METHODS

We used archival data from Travel & Group charts(T&Gs) recorded throughout the year 1973 at the Gombe
Stream Research Centre, mostly by Caroline Tutin (COPS
study) and McGrew (KIDS and TERM studies). T&Gs
were records of focal-subject samples of independently
moving chimpanzee subjects, followed by a researcher
and field assistant for variable periods, up to a whole active day (i.e. nest-to-nest). For details and results from thisdata instrument, see Goodall 1986.
For this analysis, we focussed on the T&G column
labelled FOOD, in which duration of bouts of feeding (tothe nearest 5-min) and species fed-upon by the focal subject were the only data available. Feeding was defined as
items being inserted into the mouth but not expelled, i.e.
were ingested. A feeding bout was defined as a continuous session of feeding on one prey item by the subject,from start to finish.
We also used data from 10 of the 19 T&G columns
labelled by the two-letter initials of adult and subadultmales (EV, FB, FG, GB, HG, HM, JJ, MK, SH, ST), allmembers of the Kasekela community see Table 1). The
other 9 males, mostly from the less-studied Kahama community, had too few records. A focal subject could haveup to 41 companions in a party; younger individuals, typically offspring, were not included, as they did not moveindependently. These data yielded number of companionswith the focal subject in 5-min, one-zero frequency units.
Number of companions was preferred as the dependent
variable based on semantics (i.e. party is a plural term, so
a party-size of one is nonsensical) and computation (i.e.
allows for a party size of zero to be a lone individual).
Finally, we noted the column labelled IN CA, which
distinguished In Camp versus Out of Camp records.
(Camp denoted the feeding area, where provisioning withbananas was done.) The 50 In Camp records found were
excluded from further analyses, as being unnatural.
The following data were extracted: Number of companions at the beginning and end of each Out of Campfeeding bout (n = 558), plus party size 30 min before and30 min after the feeding bout (n = 69). Thirty min was arbitrarily chosen, based on comparing behaviour ‘beforeduring-after’ feeding, in earlier studies of chimpanzee
behaviour (e.g. de Waal 1987). Thus data were collected atfour sequential points in time. However, we excluded allcases in which the ‘before’ or ‘after’ data were compromised by another feeding bout during the intervening 30min interval, which greatly reduced the number of fourpoint sequences.
Data were entered into a Microsoft Access 2013 database (spreadsheet). Statistical testing was non-parametric,
as the conditions for parametric testing could not be met(Siegel & Castellan 1988). Accordingly, we used mediansrather than means, and alpha was set at 0.05, one-tailed.

RESULTS

Number of companions varied across the four conditions (stages): Friedman two-way analysis of variance, n= 10, k = 4, χ2 = 70.8, p < 0.001 (see Table 1, Figure 1).
Pair-wise comparisons between conditions also varied inthree of four cases: Number of companions decreased in

Figure 1. Number of companions (median of medians) for
10 independently-ranging male chimpanzees
at Gombe, before, during and after bouts offeeding.

number from 30 min before the start of the next feeding
bout (Binomial test, n = 8, excluding ties, p = 0.035) andincreased in number 30 min after the end of the last feeding (n = 5, p = 0.01). Despite having equal medians, number of companions increased from the start to the end of
feeding bouts (n = 10, p = 0.03). There was no differencein number of companions between 30 min before versus
30 min after feeding bouts (n =8, p > 0.05).
Thus, males fed with fewer companions than they
had a half hour before the next bout of feeding, and a halfhour after finishing feeding they had more companions.
Moreover, in a typical feeding bout, a male had morecompanions at the end of it than he had at the beginning.
Number of companions (effectively party size, as above)
before and after feeding did not differ in number.

DISCUSSION

The hypothesis was upheld. Gombe’s adult malechimpanzees somehow manage their fission-fusioning,
so that they were in smaller parties when feeding versus
when not feeding. Thus, sequential, individual data at themore specific, proximate level agree with correlational
data at the more general, ultimate level. The apes mayachieve this by avoiding others before starting to feed,
only then to seek others after feeding. However, this dataset yields no details on how the increases and decreases in
number of companions are managed. Explanation awaits
further detailed ethological study.
We made no prediction about whether party size
would increase or decrease during an individual’s feeding
bout, as either alternative is a reasonable expectation. A
male might recruit others (e.g. Clark & Wrangham 1993)
and so increase party size, or party sizes might shrink as
the patch’s food is depleted. Or, party size might remain
constant over a feeding bout, as on average, differing
tendencies counter-balance. In any case, Gombe’s males
had slightly more companions at the end of a feeding boutthan at its outset. This could indicate recruitment or justrandom but cumulative discovery of the food source byothers. Again, to explain this finding requires more focussed, specific data than are available here.

ACKNOWLEDGEMENTS

We thank: Gombe colleagues, and especially Caroline

22
Tutin, for retrospective use of their data; Miami University
Honors Program and Linda Marchant for funding for AB; The
Leverhulme Trust for funding for WM; Noriko Itoh for comments on the manuscript.

REFERENCES

Aureli F, Schaffner CM, Boesch C, Bearder SK, Call J,
Chapman CA, Connor R, Di Fiore A, Dunbar RIM,
Henzi SP, Holekamp K, Korstjens AH, Layton R, Lee P,
Lehmann J, Manson JH, Ramos Fernandez G, Strier KB,
van Schaik CP 2008. Fission-fusion dynamics: new research frameworks. Cur Anthropol 49:627–654.
Clark AP, Wrangham RW 1993. Acoustic analysis of wild
chimpanzee pant hoots: do Kibale Forest chimpanzees
have an acoustically different food arrival pant hoot? Am J
Primatol 31:99–109.
Goodall JvL 1968. The behaviour of free-living chimpanzees in the Gombe Stream Reserve. Anim Behav Monog
1:161–311.
Goodall J 1986. The Chimpanzees of Gombe. Harvard
University Press, Oxford.
Hashimoto C, Suzuki S, Takenoshita Y, Yamagiwa J,
Basebose AK, Furuichi T 2003. How fruit abundance affects the chimpanzee party size: a comparison between
four study sites. Primates 44:77–81.
Isabirye-Basuta G 1988. Food competition among individuals
in a fre-ranging chimpanzee community in Kibale Forest,
Uganda. Behaviour 105:135–147.
Itoh N, Nishida T 2007. Chimpanzee grouping patterns andfood availability in Mahale Mountains National Park,
Tanzania. Primates 48:87–96.
Kummer H 2008. Comment on Aurelli et al.. Cur Anthropol
49:644–645.
Matsumoto-Oda A, Hosaka K, Huffman MA, Kawanaka K
1998. Factors affecting party size in chimpanzees of theMahale Mountains. Int J Primatol 19:999–1011.
Newton-Fisher NE 1999. Association by male chimpanzees:
a social tactic? Behaviour 136:705–730.
Newton-Fisher NE, Reynolds V, Plumptre A 2000. Food supply and chimpanzee (Pan troglodytes schweinfurthii) party
size in the Budongo Forest Reserve, Uganda. Int J Primatol
21:613–628.
Nishida T 1968. The social group of wild chimpanzees in the
Mahali Mountains. Primates 9:167–224.
Siegel S, Castellan, NJ 1988. Nonparametric Statistics for the
Behavioral Sciences. McGraw-Hill, New York.
Sugiyama Y 1968. Social organization of chimpanzees in the
Budongo Forest, Uganda. Primates 9:225–258.
de Waal FBM 1987. Tension regulation and nonreproductivefunctions of sex in captive bonobos (Pan paniscus). Natl
Geog Res 3:318–335.
Wrangham RW 1977. Feeding behaviour of chimpanzees inGombe National Park. In: Primate Ecology. Clutton-Brock
TH (ed), Academic Press, London, pp. 504–538.
Yamagiwa J 2008. Comment on Aurelli et al.. Cur Anthropol
49:645–646.

Pan Africa News, 20(2), December 2013

<NOTE>A Juvenile Chimpanzee Playedwith a Live Moth
Michio Nakamura
Wildlife Research Center, Kyoto University, Japan(E-mail: nakamura@wrc.kyoto-u.ac.jp)

Introduction

Although it is not common for chimpanzees (Pantroglodytes) to capture and toy with small mammals or
birds without eating them, several reports have indicatedthat this does sometimes occur (e.g., Hirata et al. 2001;
Zamma 2002; Carvalho et al. 2010). These reports compared such incidents with chimpanzees’ hunting behaviors because it seems puzzling that the chimpanzees did
not eat the meat. However, it is possible that a chimpanzee
may show an interest in a living organism for its animacy
per se, with no intention of eating it.
This paper reports the case of a juvenile female chimpanzee at Mahale that captured and played with a live
moth (Lepidoptera). Although chimpanzees frequently
prey on several insect species (mostly social insects)(e.g., McGrew 1992; Fuse 2013), Mahale chimpanzees eat
moths only rarely (Nishida & Uehara 1983).

OBSERVATIONThe observation focused on chimpanzees in the
M group at the Mahale Mountains National Park (seeNishida 2012 for details of the study site). At 09:19 a.m.
on 12 October 2004, two adult female chimpanzees andtheir offspring had been taking a rest under a dense bushsince 08:54 a.m. when Ichiro (IH: a 1-year-old male) emitted a slight huu call and was observed to be watchingsomething on the ground. When IH touched the object
with his finger, it moved, and I could then see that it wasa large moth, about 5 – 6 cm in size and beige in color. Its
body was thick, and its forewings were much longer thanits hindwings. IH’s mother, Ikocha (IK), immediately approached him, put him on her belly, moved away, and sat
about 1 m away from the moth. Soon, IH’s older sister,
Imani (IM: a 6-year-old female), went to where the mothlay and started to investigate it. Because my initial view
was largely obstructed by dense bush, I moved to the side
to gain a better view and started to take a video.
IM captured the moth without killing it and started
to play with it, sometimes vigorously, sometimes calmly.
Given her occasional play face and play pant, IM’s series of behaviors was regarded as playful. I summarize
the behavioral patterns observed during IM’s play inTable 1 (see Video 1: available online at mahale.main.jp/
PAN/20_2/20(2)_03 .html). The moth sometimes fluttered
its wings while being pinched between IM’s fingers or
lips. Although IM frequently put the moth in her lips or
touched it with her lip, she did not try to eat it.
While IM was playing with the moth, IH approached
her twice, and she responded playfully to him. At 09:25,
IM put the moth in her groin pocket and slowly walked
away into the bush; at 09:26, I lost sight of her because I
could go no further into the bush. When she was observedPan Africa News, 20(2), December 201323

again 4 min later, she no longer had the moth

Discussion

Unfortunately, I could not collect the moth or identify the species. However, from its appearance andsize, I suspect that it was probably a hawk moth (family
Sphingidae). Although Nishida & Uehara (1983) listed
unidentified moths (larvae and imagoes) in the food list of
Mahale chimpanzees, with no description as to how they
ate them, IM’s behaviors described here did not look likea feeding attempt.
The total observation time during which IH and IM
showed interest in and interacted with the moth was only
about 7 min, but there were some similarities between
their behaviors and chimpanzees’ play with small vertebrates. For example, as Zamma (2002) reported in a caseof a chimpanzee playing with a live squirrel, it seemedthat IM too was enjoying the response of the moth, whichsometimes fluttered. When the moth was motionless, shesometimes put her lower lip on it as if she were trying
to activate the moth’s movement again. She was careful
enough in her handling of the moth not to flatten it, but
because she finally tore off a part of a wing, it is difficult
to say that she had no intention of killing the moth.
Unfortunately, I did not observe what finally happened to the moth, but since the moth had already become
weak, I suspect that it soon became inactive, and thus IM
might have lost interest and abandoned it. This differsfrom former observations of mammalian and avian play
toys, in which the interest of the chimpanzees was main-

Figure 1. IM putting a forewing of the moth between her
lips while lying prone (image taken from video
footage).

tained for much longer, and they sometimes groomed ortreated the animals like dolls even after the animals weredead (Hirata et al. 2001; Zamma 2002; Carvalho et al.
2010). Such differences may be due to differences in thesizes and shapes of the target animals.
Interestingly, although it was the infant, IH, who first
showed interest in the moth, his mother immediately took
him away from the insect. Some insects are poisonous or
spiny, so the mother may have been protecting her young
son from such potential dangers.

Table 1. IM’s behaviors toward the moth in relation to her body postures.

Behavior towards the mothBody posture

Puts the moth on the ground and hits it with her hands

Sitting

Throws the moth with one handStanding on all fours

Puts a fore wing of the moth between her lips (Figure 1),
sometimes shaking her head

Sitting, lying supine, lying prone, rolling sideways, turning
head over heels

Grabs the moth together with fallen leaves and lays her bodyon a pile of leaves

Sitting, lying prone

Pinches the moth with her hand or foot and holds it up in the
air

Lying supine

Pinches the moth with her hand and brings it close to her
nose or mouth

Sitting, lying supine

Kicks the moth backwards with her footStanding on all fours

Puts the moth in her groin pocket

Lying supine

Walks with the moth in her groin pocketStanding on all fours

Pinches the moth’s forewings with both hands and pulls
slowly, tearing one wing (Figure 2)

Lying supine

Puts the moth between her arm and the side of her body

Lying on her side

Touches the moth with her footStanding on all fours

24

Pan Africa News, 20(2), December 2013

<NEWS>

The 1st MWCS ScholarshipStudent Awarded Diploma
Kazuhiko Hosaka
C o - c h a i r m a n , Ma h ale Wil dlife C o n se r v a t i o n S o c ie t y(MWCS)(E-mail: khosaka1@gmail.com)

Figure 2. IM pinching both wings of the moth with both
hands and slowly pulling it apart on the solesof her feet while lying supine (image taken fromvideo footage).

It is common for human children to capture, playwith, and sometimes kill still-living insects with no apparent purpose. It is possible that both humans and chimpanzees, especially youngsters, share a common inclination to become interested in, and sometimes enjoy playing
with, an animate object.

Acknowledgements

I thank COSTECH, TAWIRI, and TANAPA for theirpermission to conduct long-term research at Mahale. The
study was supported by Kakenhi (#19255008, #19107007, and#21770262).

References

Carvalho S, Yamanashi Y, Yamakoshi G, Matsuzawa T 2010.
Bird in the hand: Bossou chimpanzees (Pan troglodytes)
capture West African wood-owls (Ciccaba woodfordi) but
not to eat. Pan Afr News 17:6–9.
Fuse M 2013. Chimpanzees detect ant-inhabited dead branches
and stems: a study of the utilization of plant–ant relationships
in the Mahale Mountains, Tanzania. Primates 54:385–397.
Hirata S, Yamakoshi G, Fujita S, Ohashi G, Matsuzawa T 2001.
Capturing and toying with hyraxes (Dendrohyrax dorsalis)by wild chimpanzees (Pan troglodytes) at Bossou, Guinea.
Am J Primatol 53:93–97.
McGrew WC 1992. Chimpanzee Material Culture: Implicationsfor Human Evolution. Cambridge University Press,
Cambridge.
Nishida T 2012. Chimpanzees of the Lakeshore: Natural
History and Culture at Mahale. Cambridge University Press,
Cambridge.
Nishida T, Uehara S 1983. Natural diet of chimpanzees (Pantroglodytes schweinfurthii): Long-term record from the
Mahale Mountains, Tanzania. Afr Study Monogr 3:109–130.
Zamma K 2002. A chimpanzee trifling with a squirrel: pleasure
derived from teasing. Pan Afr News 9:9–11.

I am pleased to announce that Mr. Butati R. Nyundo,
a grantee of the MWCS Special Scholarship, successfullycompleted a two-year course programme for ordinary
diploma in wildlife management at the College of African
Wildlife Management, Mweka (CAWM), Tanzania andattended the graduation ceremony taking place on the
25th of October, 2013 in order to get his diploma (Photo1). This was MWCS’s first attempt to offer an opportunity
for high education to a prospective young person eager
for wildlife conservation at Mahale. We will examine
how it will work and what we can do to contribute to thecommunity-based conservation in this area, while keeping in touch with Mr. Nyundo and giving him support, if
necessary. We hope conservational efforts will thrive at
Mahale in the hands of the Tongwe, the indigenous people
there. Mr. Nyundo himself aspires to work for the Mahale
Mountains National Park and to contribute to protectionof the natural environments from any kind of harmful
factors.
The following is his report on his three-month activities as a volunteer teacher at Buhingu Secondary School
and Katumbi Primary School. Such activities had been
assigned him by MWCS as an extended programme of thescholarship. Acknowledgements are due to all those whohave supported our aims and activities as members and/or
donors.

Photo 1. Mr. Butati R. Nyundo after graduation ceremony.

My teaching experience at myhometown
Butati R. Nyundo
Mahale Mountains National Park

As soon as I completed my studies at CAWM, I wentPan Africa News, 20(2), December 2013
back to my hometown Kigoma for volunteer activities. It
was a welcoming idea to me as I had desired to have sucha chance as to contribute something to my community
consisting mainly of the Tongwe people. Teaching secondary school students and primary school kids turned
out an incredible experience.
I started to teach as a volunteer at Buhingu Secondary
School in the middle of January 2013. I considered it
particularly valuable to work for the secondary school,
because it enrolled students from as many as six villages,
which meant that I could be involved in the biggest part ofthe community. On the first day, I was introduced to theschool administration and the students with a warm welcome from everyone.
I was assigned to teach biology to all classes of Form
1–4 and geography to one class of Form 3. Teachers andstudents were very friendly to me, and it did not take meso long to get used to the environment.
Students looked inspired by my presence as theycame to know that a person like me from their own community could attain higher education. And I in turn was
motivated and encouraged to work harder by their eagerness to learn new ideas especially concerning with wildlife and environmental conservation. They made efforts
to improve their environment by planting trees aroundthe school compound and did their best to stop bush fires.
I gave several talks about environmental conservation todifferent classes and the Roots and Shoots Tanzania club.
The school management acknowledged my efforts
and influence on the students and appointed me as assistant of sports and counseling master. I took the school
football team to three different games where we got two
wins and a draw. I also made several advices to students
how they should study to be more successful. How wonderful it was for me to see boys and girls work harder as aresult of my advices!
However, due to remoteness, the school was facing
many problems concerning its facilities and equipment.
I interviewed with the staff and made a list of the prioritized needs at Buhingu Secondary School, which wasthen attached to this report submitted to MWCS.
In early March, Nomad Tanzania, a safari company
managing a camp at Kangwena Beach of the Mahale
Mountains National Park, offered us a day trip to the parkby its boat. The trip comprised of eighteen students, one
teacher and me. We took students of thirteen years or
more, in accordance with the code of conduct for chimpanzee viewing. My role was to accompany them as their
guide, wildlife expert and a native of that area. I did mybest to offer the students the explanation of the park; thehistory of Mahale, conservation activities, resources,
chimpanzees, Lake Tanganyika and cichlids uniqueto this lake (Photo 2). We had a chance to meet Mr.
Shunkichi Hanamura of Kyoto University, who stayed at
the research station as a member of the Mahale Mountains
Chimpanzee Research Project that had persisted since1965. The students enjoyed the trip very much and seemed
to comprehend the significance of conservation activities
undertaken in the area.
I spent the final month of my volunteer work in teaching at Katumbi Primary School, which was very special

25
to me because Katumbi is my home village. Children start
to attend school at the age of 5 or 6 years. Training suchyoung kids requires more effort than at secondary education, but unfortunately, the school had a severe shortageof teachers. I was assigned to teach science and geography
to Class 7, 6, 5 and 4. Besides a shortage of teachers, theschool was struggling with the poor conditions concerning desks, sports equipment and some facilities in need ofrepair. Nevertheless, some students performed very well
above average in their studies.
Now that three month volunteer activities have been
over, I was very happy and proud to have such an opportunity as to work at Buhingu Secondary and Katumbi
Primary Schools. Once again, I would like to thank
MWCS for their full support of my two years diploma
course at the CAWM and the three months livelihood support during my volunteer activities.

Photo 2. Mr. Nyundo answering questions from secondary
school students in the forest during a day trip
to Mahale Mts. National Park. Photo by Mr.
Shunkichi Hanamura.

<BOOK INFO>

Primates and Cetaceans: Field
Research and Conservation ofComplex Mammalian Societies
Edited by Juichi Yamagiwa andLeszek Karczmarski
In this book, the editors present a view of the socioecology of primates and cetaceans in a comparative perspective to elucidate the social evolution of highly intellectual
mammals in terrestrial and aquatic environments. Despite
obvious differences in morphology and eco-physiology,
there are many cases of comparable, sometimes strikingly
similar patterns of sociobehavioral complexity. A numberof long-term field studies have accumulated a substantial
amount of data on the life history of various taxa, foraging ecology, social and sexual relationships, demography,
and various patterns of behavior: from dynamic fission–Pan Africa News, 20(2), December 2013
fusion to long-term stable societies; from male-bonded tobisexually-bonded to matrilineal groups.
Primatologists and cetologists have come together
to provide four evolutionary themes: (1) social complexity and behavioral plasticity, (2) life history strategies
and social evolution, (3) the interface between behavior,
demography, and conservation, and (4) selected topics incomparative behavior. These comparisons of taxa that are
evolutionarily distant but live in comparable complex sociocognitive environments boost our appreciation of their
sophisticated mammalian societies and can advance ourunderstanding of the ecological factors that have shaped
their social evolution. This knowledge also facilitates a
better understanding of the day-to-day challenges theseanimals face in the human-dominated world and may improve the capacity and effectiveness of our conservation
efforts.

439 pages
Springer
Publication year: 2014
Hardcover 149,99 €; ISBN 978-4-431-54522-4
eBook 124,94 €; ISBN 978-4-431-54523-1
Contents
Part 1: Social Ecology
1. How ecological conditions affect the abundance andsocial organization of folivorous monkeys / Colin A.
Chapman, Tamaini V. Snaith & Jan F. Gogarten
2. Dusky dolphins: Flexibility in foraging and social
strategies / Bernd Würsig & Heidi C. Pearson
3. Socioecological f lexibility of gorillas and chimpanzees / Juichi Yamagiwa & Augustin Kanyunyi
Basabose
4. You are what you eat: Foraging specializations andtheir influence on the social organization and behaviour of killer whales / John K. B. Ford & Graeme M.
Ellis
5. Japanese macaques: Habitat-driven divergence in social dynamics / Goro Hanya
6. Shark Bay bottlenose dolphins: A case study for defining and measuring sociality / Margaret A. Stanton& Janet Mann

26
Part 2: Life History and Social Evolution
7. Female coexistence and competition in ringtailed
lemurs: A review of a long-term study at Berenty,
Madagascar / Yukio Takahata, Naoki Koyama,
Shin’ichiro Ichino, Naomi Miyamoto & Takayo Soma
8. Social structure and life history of bottlenose dolphins near Sarasota Bay, Florida: Insights from four
decades and five generations / Randall S. Wells
9. Life history tactics in monkeys and apes: Focus on
female dispersal species / Juichi Yamagiwa, Yukiko
Shimooka & David S. Sprague
10. Social conflict management in primates: Is there acase for dolphins? / Marina Cords & Janet Mann
11. Evolution of small-group territoriality in gibbons /Warren Y. Brockelman, Anuttara Nathalang, David B.
Greenberg & Udomlux Suwanvecho
Par t 3: Demogra phy, Gene t ic s , and Is sue s inConservation
12. Northern muriqui monkeys: Behavior, demography,
and conservation / Karen B. Strier
13. Indo-Pacific humpback dolphins: A demographic
perspective of a threatened species / Shiang-Lin
Huang & Leszek Karczmarski
14. Mountain gorillas: A shifting demographic landscape / Elizabeth A. Williamson
15. Population genetics in the conservation of cetaceans
and primates / Kimberly Andrews
16. Eco-toxicants: A growing global threat / Victoria
Tornero, Teresa J. Sylvina, Randall S. Wells &Jatinder Singh
Part 4: Selected Topics in Comparative Behavior
17. Observing and quantifying cetacean behavior in thewild: Current problems, limitations and future directions / Janet Mann & Bernd Würsig
18. Social network analysis: Applications to primate
and cetacean societies / Margaret A. Stanton & Janet
Mann
19. Social touch in apes and dolphins / Michio
Nakamura & Mai Sakai
20. Non-conceptive sexual interactions in monkeys,
apes, and toothed whales / Takeshi Furuichi, Richard
Connor & Chie Hashimoto
21. A mix of species: Associations of heterospecifics among primates and dolphins / Marina Cords &Bernd WürsigPan Africa News, Vol. 20, No.2
Published in December, 2013
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)