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

JUNE 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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Pan Africa News publishes articles, notes, reviews, forums,
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<ARTICLE>

The Population and Habitat
Preferences of Chimpanzees in
Non-Protected Areas of Tanzania
Hideshi Ogawa1, Midori Yoshikawa2& Gen’ichi Idani3
1 School of International Liberal Studies, Chukyo University
2 United Graduate School of Agricultural Science,
Tokyo University of Agriculture and Technology
3 Wildlife Research Center, Kyoto University(E-mail: hogawa@lets.chukyo-u.ac.jp)

VOL. 20, NO. 1

Contents<ARTICLE>
The Population and Habitat Preferences of
Chimpanzees in Non-Protected Areas of Tanzania
Hideshi Ogawa, Midori Yoshikawa &Gen’ichi Idani				1<NOTE>
Well Digging by Semliki Chimpanzees: New Data on
Laterality and Possible Significance of Hydrology
William C. McGrew, Linda F. Marchant, Charlotte
L.R. Payne, Timothy H. Webster & Kevin D. Hunt						 5<NOTE>
Newly Observed Bird Consumption by a Chimpanzee(Pan troglodytes) at Bossou, Guinea, West Africa
Lira Yu, Boniface Zogbila & Tetsuro Matsuzawa 8<NEWS>The Process of Creation of a New Protected Area in the
Democratic Republic of Congo: The Case of the Iyondji
Community Bonobo Reserve
Jef Dupain, Andrew Fowler, Phila Kasalevo,
Tetsuya Sakamaki, Bongoli Lingomo, Theo Way,David Williams, Takeshi Furuichi &Charly Facheux 			 10<BOOK INFO>Groups: The Evolution of Human Sociality		

13<FROM THE EDITORS>			

13

INTRODUCTION

Tanzania is the most eastern region of the distributionof eastern chimpanzees (Pan troglodytes schweinfurthii)(Kano 1972). According to Kano’s survey in 1965–67(Kano 1972), chimpanzees inhabited the eastern shore of
Lake Tanganyika: a) Gombe (Gombe National Park), b)
Lilanshimba, c) Ugalla, d) Masito, e) Mukuyu, f) Mahale(Mahale Mountains National Park), g) Karobwa, and h)
Wansisi. In addition, Ogawa and others (1997) found themost southern chimpanzee population in i) Lwazi in 1996
(Figure 1). The geographical distribution of chimpanzee
beds (Yoshikawa et al. 2008) and DNA variation (Inoue

1

Pan Africa News, 20(1), June 20132
et al. 2013) indicate that the chimpanzee population iscontinuous in c–h areas. This means that there are four
local populations of chimpanzees in Tanzania: a) Gombe,b) Lilanshimba, so called “Greater Mahale” composed ofc–h areas, and i) Lwazi, as well as an introduced population on Rubondo Island in Lake Victoria. Although chimpanzees still inhabit all areas in which they were foundin 1960s (Kano 1972; Ogawa et al. 1997, 2004, 2006a,
2006b, 2007; Hernandez-Aguilar 2006; Moyer et al. 2006;
Yoshikawa et al. 2008; Ndimuligo 2008; Plumptre et al.
2010), the population density of chimpanzees in nonprotected areas has been significantly reduced from 1960s
to 2000s (Yoshikawa et al. 2008). Proper conservational
planning is essential to protect chimpanzees (Plumptre etal. 2010). However, chimpanzees in such areas have beenmuch less studied than those in two national parks, a)Gombe and f) Mahale. Therefore, in addition to our extensive field surveys in 1994–2008 (Ogawa et al. 1997, 2004,
2006a, 2006b, 2007; Yoshikawa et al. 2008) and estimation of population densities of chimpanzees (Yoshikawa
et al. 2008), we conducted further surveys in 2010–2011
and GIS analyses of chimpanzee habitats in non-protectedareas of Tanzania.

100kma)GombeKigomab)Lilanshimbad)Masitoe)Mukuyu

to Tabora 05°

railway

Uvinzac)Ugalla
200kmg)Ka j) Mishamok) Sabagag)Karobwal) Niamansi

Gombe
Mahalef)Mahale5°

Dar es Salaam

Tanzania

Mpanda10°

Lake Tanganyikah)Wansisi

30°

35°

07°

)

Sumbawangai)Lwazi
Ufipa Escarpment

From 1994 to 2011, we conducted transect bed censuses in all chimpanzee habitats out of national parks:b, c, d, e, g, h, and i, (Ogawa et al. 1997, 2004, 2006a,
2006b, 2007; Yoshikawa et al. 2008) (Figure 1, Table 1).
As a control, we conducted bed censuses at j) Mishamo,k) Sabaga, and l) Niamansi in non-habitats of chimpanzees (Figure 1, Table 1). We usually made a round trip
from our base camp along the river to the top of a hill
and walked on not only traditional paths, but also pathsof wild animals and off-path. When we found a chimpanzee bed, we recorded location of the bed using GPS,vegetation of the bed site, and perpendicular distance
between the bed and the census line. We chose the census
lines whose tracks had been recorded by GPS and/or on
1/50,000 maps, and plotted the census lines on a vegetation map using ERDAS Imagine (Leica Geosystems GIS& Mapping) based on satellite images (ETM+). We thendefined a census belt of 60 m width centered on the census line. This width was used, because the detection rateof a chimpanzee bed was constant within 30 m from thecensus lines. We further divided all of the census belts
and defined 60 × 60-m quadrats along the census line. We
then calculated (1) proportion of the area covered by evergreen forests, (2) average ground slope, and (3) average
altitude in each quadrat using a 90 m SRTM digital elevation model. Using ArcGIS (ArcView 9.3.1) (ESRI), we
calculated the distance between each bed cluster and thenearest evergreen forest with area ≥ 1 ha, and the distance
between each study site and the nearest edge of the nearest human settlement with area approximately ≥ 10 ha.

RESULTSFor GIS analyses, we used a total of 1,026 km census
lines in chimpanzee habitats and 46 km in non-habitats,yielding 17,845 60 × 60-m quadrats along the census lines(Table 1). The area was occupied by savanna woodlands

08°

Kasanga
Habitats of chimpanzees
Beds and other traces of chimpanzees

METHODS

06°

Tunduma
30°

31°

09°

32°

Figure 1. Distribution of chimpanzees in Tanzania.

(86.9%), evergreen forests (5.9%), and others such as
grasslands, bamboo thickets, cultivated fields, and humansettlements (7.2%).
In chimpanzee habitats, there were 665 beds in 186
quadrats, while there were no beds in 17,659 quadrats.
Evergreen forests occupied more area, 25.0 ± 35.5% (mean± SD) (range: 0–100), in the quadrats with beds than in
the quadrats without beds, 5.7 ± 18.8% (0–100) (Figure2(1), Mann-Whitney U test, n1 = 186, n2 = 17,659, z = 12.6,
p < 0.05). The slope of the ground was steeper, 10.1 ± 7.1°
(0.6–30.8), in the quadrats with beds than the quadrats
without beds, 6.5 ± 22.8° (0.1–37.7) (Figure 2(2), z = 8.2,
p < 0.05). The difference was not significant between
the altitude in the quadrats with beds, 1,217 ± 154.8 m(797–1,814), and those in the quadrats without beds, 1,216± 182.0 m (780–1,876) (Figure 2(3), z = 1.5, ns.).
The ground slope and altitude of all quadrats withbeds were between 0.6° and 30.8°, and between 797 m
and 1,814 m. If we regard such quadrats as possible bed
sites of chimpanzees, the possible bed sites were 97.4% in
chimpanzee habitats and 100% in non-habitats (Table 1).
We recorded a total of 2,154 beds within and out of
30m from the census lines. Chimpanzees made 1,085
beds (50.4%) in forests, 1,020 beds (47.3%) in woodlands,
and 49 beds (2.3%) in mixed vegetations (Table 1). Beds
in woodlands were 0.15 km (0.01–0.64) from the nearest
forest (Table 1).
The population density of chimpanzees was positively
correlated with the proportion of forest in the area (Table
1, Kendall rank correlation test, n = 10: b-e, g-l areas, τ =0.58, z = 2.3, p < 0.05), and with the distance between thearea and the nearest human settlement (Table 1, τ = 0.71, z= 2.9, p < 0.05).

Location

05°43'-06°30'S Sep '94 6
29°45'-30°13'E

0

-

-

71
(102)
26
(52)
95
(154)
17
(78)
42
(69)
665
(2,154)

35
(187)
379
(1,512)

-

-

6.2

2.9

0.3

16.9

7.9

8.7

4.1

7.6

-

-

-

5.4
1,072
(0.2–31.5) (795–1,315)
7.8
890(0.1–25.9) (780–1,027)
8.1
1,241
(0.2–37.7) (967–1,876)
7.2
1,422
(0.5–22.1) (1,178–1,823)
6.8
1,349
(0.2–29.1) (780–1,889)
6.2
1,211
(0.1–37.7) (780–1,889)

5.6
1,115
(0.5–29.1) (911–1,302)
5.7
1,210
(0.1–33.5) (1,041–1,621)

-

-

97.4

94.5

99.2

97.5

89.3

99.1

97.7

99.7

Altitude (m)2 Proportaionof possible
bed site
(%)3

-

-

39
(38.2%)
3(5.8%)
47(30.5%)
0(0.0%)
27(39.1%)
1,085(50.4%)

31(16.6%)
938(62.0%)

Bed site
Numberof beds inevergreen
forest4

Total
Non-habitatj) Mishamo, k) Sabaga, l) Niamansi7 Aug '03, Sep '05,
46
0
0.5
3.4
1,357
100.0
Aug '06(0.2–19.1) (1,105–1,602)
1. Numbers of beds within 30 m from the census line are shown. Total numbers of beds found in the areas are shown in parentheses.
2. Mean numbers in each 60 × 60-m quadrat are shown. Ranges are shown in parentheses.
3. Percentages of the quadrats whose ground slope was between 0.6°and 30.8°, and whose altitude was between 797 m and 1,814 m are shown.
4. % = (Number of beds in forests) / (Number of all beds) × 100.
5. Densities of individuals excluding immature sleeping with their mother are shown with 95% confidence intervals in parentheses (Yoshikawa et al. 2008).
6. We did not conduct a bed census, but stayed during the periods in the areas: a and f.
7. The location was 05°38'S, 30°28'E at j, 05°58'S, 30°45'E at k, and 05°56'S, 30°57'E at l.

f) Mahale

0

1,026

138

47

125

31

78

488

119

Bed census
Study periods Census Number Proportion Groundslope (°)2
distance of beds1ofevergreen(km)
forest (%)2

HabitatNon-protected areab) Lilanshimba
05°07'-05°14'S Jul '01, Aug '06
30°07'-30°14'Ec) Ugalla
05°09'-05°52'S Sep '95, Aug '96,
30°23'-31°01'E Aug '97, Jul '01,
Feb '03, Jul '03,
Aug '07, Aug '08,
Dec '10-Mar '11d) Masito
05°10'-05°35'S Aug '03, Aug '05
29°50'-30°35'Ee) Mukuyu
05°22'-05°51'S Aug '05
29°44'-30°07'Eg) Karobwa
05°51'-06°10'S Aug '03
29°55'-30°31'Eh) Wansisi
06°19'-06°38'S Aug '01
30°27'-30°43'Ei) Lwazi
07°48'-08°28'S Aug '01, Jul '03,
30°55'-31°10'E Aug '05
Subtotal
05°07'-08°28'S
29°44'-31°01'E
Protected area (national park)a) Gombe
04°37'-04°45'S Sep '03, Aug '08 6
29°36'-29°40'E

Area

Table 1. Environments of chimpanzee habitats in Tanzania.

-

-

-

0.19
(0.04–0.51)
0.08(0.01–0.24)
0.10(0.01–0.36)
0.16(0.08–0.36)
0.14(0.01–0.43)
0.15(0.01–0.64)

0.13(0.01–0.64)
0.15(0.01–0.62)

6.8

-

-

20.3

11.7

13.6

15.5

8.3

16.5

32.5

14.4

0.00

-

-

0.12
(0.04–0.49)
0.05(0.02–0.08)
0.03(0.02–0.08)
0.02(0.02–0.08)
0.01(0.00–0.03)
0.05

0.02
(0.01–0.05)
0.10(0.07–0.18)

Distance
Density ofDistancefromto human chimpanzees5
woodland settlement (individuals/
beds to(km)
km2)
evergreen
forest (km)-11,092

1,600

52

9,440

1,250

500

900

1,100

2,200

3,350

140

Habitat
range
(km2)

0

approx 1,300

approx 500

approx 100

264(88–1,078)
55(22–198)29(18–72)10(5–40)
18(0–38)
714

3
(1–7)
335
(235–603)

this study

Moyer et al. 2006;
Ndimuligo 2008;
Plumptre et al. 2010
Kano 1972; Moyer et
al. 2006; Ndimuligo
2008

Ogawa et al. 1997;
Yoshikawa et al. 2008

Ogawa et al. 2004;
Yoshikawa et al. 2008

Ogawa et al. 2006a;
Yoshikawa et al. 2008

Yoshikawa et al. 2008

Yoshikawa et al. 2008

Hernandez-Aguilar
2006; Ogawa et al.
2007; Yoshikawa et al.
2008; this study

Ogawa et al. 2006b;
Yoshikawa et al. 2008

Number of Reference
chimpanzees(individuals)

Pan Africa News, 20(1), June 20133

Pan Africa News, 20(1), June 2013

4(1) Proportion of evergreen forest (%)(2) Slope of the ground (°)(3) Altitude (m)Figure 2. Conditions in quadrats containing chimpanzee beds in Tanzania.
(1) Percentage of evergreen forest area, (2) slope of the ground, and (3) altitude are compared between 60×60-m quadrats
containing chimpanzee beds and quadrats without beds. With beds: quadrats containing beds. Without beds: quadrats
without beds. Area: b=Lilanshimba, c=Ugalla, d=Masito, e=Mukuyu, g=Karobwa, h=Wansisi, i=Lwazi, N=non-habitat: j)
Mishamo, k) Sabaga, and l) Niamansi.

DISCUSSION

Most of chimpanzee habitats in Tanzania are savanna
woodlands (locally called “miombo” woodlands) dominated by deciduous trees of Brachystegia and Julbernardia
with small proportion of evergreen riverine forests (Kano
1972). However, 47.3% of beds were in forests and all
beds were within 0.64 km from a forest. In addition, thequadrats with chimpanzee beds had a greater proportion of forest than the quadrats without beds. Evergreen
forests are indispensable for chimpanzees to make beds
especially during the dry season, when most woodlandtrees lose their leaves (Hernandez-Aguilar 2006; Ogawa
et al. 2007). In addition, forests provide chimpanzees withsafe sleeping sites. Because forests have fewer grasses
and more trees than woodlands, carnivores such as lions(Panthera leo) and hyenas (Crocuta crocuta) are not able
to hide themselves for hunting, and chimpanzees canmore easily escape into trees (Hernandez-Aguilar 2006).
In addition to forests, hilly areas are also critical for chimpanzees in Tanzania, as they prefer making beds in steep
slopes (Kano 1972; Hernandez-Aguilar 2006; Ogawa et
al. 2007). As a result, chimpanzee habitats in Tanzania
had many evergreen forests and hilly areas.
However, the quality of chimpanzee habitats has become worse over time, caused by habitat destruction for
cultivation, firewood, charcoal, and commercial use oftimbers (Ogawa et al. 1997, 2004, 2006a, 2006b, 2007).
The population density of chimpanzees shows that the
number of chimpanzees outside national parks in Tanzania
has been reduced from approximately 1,300 (Kano 1972)
to 700 (Yoshikawa et al. 2008) during the last 40 years.
Even including 100 chimpanzees in a) Gombe and its
surrounding area (Moyer et al. 2006; Ndimuligo 2008;
Plumptre et al. 2010) and 500 chimpanzees in f) Mahale(Kano 1972; Moyer et al. 2006; Plumptre et al. 2010),
the total number of chimpanzees in Tanzania may be no
more than 1,300 individuals. Although another survey
estimated that 2,800 chimpanzees inhabited Tanzania(Moyer et al. 2006; Plumptre et al. 2010), both surveys
showed that more than half of all Tanzanian chimpanzees
inhabited non-protected areas. The estimated populationof chimpanzees in Tanzania ranges from 1,300 to 2,800.
However, similar methods of bed censuses in 1960s (Kano

1972) and 2000s (Yoshikawa et al. 2008) indicate thatthe population of chimpanzees in non-protected areas of
Tanzania has been remarkably reduced.
A huge continuous habitat, Greater Mahale, isprecious property of Tanzania. In e) Mukuyu and g)
Karobwa, however, people originally from Burundi set
many snares around the Mishamo settlement. Such poachers actually killed chimpanzees in these areas (Ogawa et
al. 2006a). In b) Lilanshimba, the Lugufu settlement for
Congolese refugees was established within chimpanzee
habitat in 1997, and cultivation and poaching by theserefugees have reduced chimpanzees (Ogawa et al. 2006b).
Small isolated populations in b) Lilanshimba and i) Lwazi
may be faced with extinction. It is urgently necessary toprotect chimpanzees in these two areas, as well as chimpanzees in Greater Mahale.
We found that the population density of chimpanzees is lower in areas close to human settlements. To
make matters worse, more and more immigrant farmers
cultivated fields except for steep hills in e) Mukuyu andg) Karobwa, and h) Wansisi (Ogawa et al. 2004, 2006a).
Such haphazard deforestation reduces chimpanzee habitats and does not necessarily benefit future generationsof Tanzanian people. It is necessary to adopt sustainable management of trees and other natural resources.
Effective conservation plans are possible, because chimpanzees prefer hilly areas which are not suitable forcultivation. Especially, c) Ugalla and d) Masito still have
habitats where only a few local people temporally visit. In
collaboration with researchers, the Tanzanian government
has to make and conduct proper conservation planning toprotect chimpanzees and their habitats in Tanzania.

ACKNOWLEDGMENTS

We are grateful to: T Kano, HY Kayumbo, J Itani,
T Nishida, J Moore, ET Massawe, T Sakamaki, A Piel, F
Stewart, A Hernandez-Aguilar, K Zamma, M Nakamura, M
Koganezawa, S Araki, D Sprague, and T Nemoto. This study
was permitted by the Tanzanian authorities (COSTECH and
TAWIRI) and financially supported by a Grand-in-Aid for
Scientific Research of MEXT, Japan (C06061064; B1257597;
A17255005; C22570223; 11J07921), Global Environment
Resea rch Fu nd F061, t he env i ron ment Resea rch a nd
Technology Development Fund (D-1007) of the Ministry of theEnvironment, Japan, the Advanced Studies on the Evolutionary

Pan Africa News, 20(1), June 2013
Origins of Human Nature (AS-HOPE), and International
Training Program (ITP-HOPE), Japan Society for the Promotionof Science.

REFERENCES

Hernandez-Aguilar A 2006. Ecological and Nesting Patternsof Chimpanzees (Pan troglodytes) in Issa, Ugalla, Western
Tanzania. Ph.D. thesis, University of Southern California.
Inoue E, Tashiro Y, Ogawa H, Inoue-Murayama M, Nishida
T, Takenaka O 2013. Gene flow and genetic diversity ofchimpanzee populations in East Africa. Primate Conserv
26:67–74.
Kano T 1972. Distribution and adaptation of the chimpanzeeon the eastern shore of Lake Tanganyika. Kyoto Univ Afr
Stud 7:37–129.
Moyer D, Plumptre AJ, Pintea L, Hernandez-Aguilar A,
Moore J, Stewart F, Davenport TRB, Piel A, Kamenya S,
Mugabe H, Mpunga N, Mwangoka M 2006. Surveys of
Chimpanzees and Other Biodiversity in Western Tanzania.
Unpublished report. Arlington, VA: United States Fish andWildlife Service (USFWS).
Ndimuligo SA 2008. Assessment of Chimpanzee (Pan troglodytes) Population and Habitat in Kwitanga Forest,
Western Tanzania. Master’s dissertation, University of
Witwatersrand, Johannesburg.
Ogawa H, Kanamori M, Mukeni SH 1997. The discovery
of chimpanzees in the Loasi River area, Tanzania: A newsouthern distribution limit. Pan Afr News 4:1–3.
Ogawa H, Moore J, Kanamori M, Kamenya S 2004. Reporton the chimpanzees of the Wansisi and Makomayo areas,
Tanzania. Pan Afr News 11:3–5.
Ogawa H, Moore J, Kamenya S 2006a. Chimpanzees in the
Ntakata and Kakungu areas, Tanzania. Primate Conserv
21:97–101.
Ogawa H, Sakamaki T, Idani G 2006b. The influence ofCongolese refugees on chimpanzees in the Lilanshimba
area, Tanzania. Pan Afr News 13:21–22.
Ogawa H, Idani G, Moore J, Pintea L, Hernandez-Aguilar A
2007. Sleeping parties and nest distribution of chimpanzees in the savanna woodland area, Ugalla, Tanzania. Int J
Primatol 28:1397–1412.
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, McLennan M, Amsini F, McNeilage
A, Makana JR, Kanamori M, Hernandez-Aguilar 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, Hashimoto C, Furuichi T, Bennett E 2010.
Eastern Chimpanzee ( Pan troglodytes schweinfurthii):
Status Survey and Conservation Action Plan 2010-2020.
ISBN978-2-8317-1246-8. IUCN, Gland, Switzerland.
Yoshikawa M, Ogawa H, Sakamaki T, Idani G 2008.
Population density of chimpanzees in Tanzania. Pan Afr
News 15:17–20.

5<NOTE>

Well Digging by Semliki
Chimpanzees: New Dataon Laterality and Possible
Significance of Hydrology
William C. McGrew1, Linda F.
Marchant2, Charlotte L.R. Payne1,
Timothy H. Webster3 & Kevin D.
Hunt4
1 Dept. of Archaeolog y & Anthropolog y, Universit y ofCambridge, UK
2 Dept. of Anthropology, Miami University, USA
3 Dept. of Anthropology, Yale University, USA
4 Dept. of Anthropology, Indiana University, USA(E-mail: wcm21@cam.ac.uk)

INTRODUCTION

The wild chimpanzees (Pan troglodytes schweinfurthii) of Toro-Semliki Wildlife Reserve, Uganda, arethe only population of wild apes yet reported to dig wells
for drinking water1. They dig holes by hand in sandy riverbeds, then drink the water collected there by mouth orwith ‘sponges’ made of leaves. We previously reported
that the excavated tailings of this digging were often ‘bilobed’ in distribution on the substrate, that is, clearly
divided into separate piles of sand to the left and rightsides of the hole (McGrew et al. 2007). Because thesepiles were symmetrical in volume, we inferred that the
chimpanzees were manually ambilateral, that is, that they
did not favour one hand or the other in digging. That is,
we inferred that had their digging been right-biassed, thenthe rightside pile of sand would have been bigger.
However, our first report was no more than a pilot
study, as it was done with only 2 days (July) of intensive
etho-archaeological data collection over less than 1 km ofriverbed, only in the dry season. Furthermore, it was donewith only a single measure, ‘volume’. Here we report a
longer study done over 6 months (May–Nov.) over morethan 5 km of riverbed, done in both wet and dry seasons.
The aim of the follow-up study was to replicate and extend the pilot study’s findings, but in the process several
unexpected factors were revealed, chiefly hydrologicalones.
In the first study in 2006, flow of surface water in the
bed of the Mugiri River had ceased, leaving only scattered, stagnant pools. In some places, the sandy substrate
was still moist, but in others it was dry. In the second
study in 2008, surface water flow never ceased, though it
varied in depth and volume, and the sand in the riverbed
was always moist. Thus, environmental affordance ofdrinking water for the chimpanzees showed marked interannual variation.
With regard to laterality, we expected (as in the pilot
study) that if the chimpanzees were manually biased at
population-level to one side or another, then overall either
left or right piles of tailings (lobes) would be larger. If
there was no bias in the pooled data, then we expected nodifference in any left- versus right-side measurements.

Pan Africa News, 20(1), June 2013

6

Figure 2. Mugiri riverbed in 2008. Two researchers
examine well on sandbar, with flowing water behind.

volume); weight (nearest 100 g, by spring balance). Allvariables except weight were for replication of 2006 data.
Note that no data could be assigned to individuals, sodata were pooled, which is usual practice for artefacts inarchaeology.

RESULTS
Figure 1. Dry Mugiri riverbed in 2006.

METHODS

Full details of the study site and methods are given
elsewhere (Hunt 2000; Hunt & McGrew 2002).
We found 121 wells, but 67 were either too disturbed
by trampling to provide data or had a single, central tailing of excavated sand, so they could not provide data onlaterality. For the remaining 54 wells that were bi-lobed(that is, had distinct left and right tailings), we recordedby tape measure (cm) the following variables: well length,
width and depth; length, width and depth of water inwell (if present); mean distance to nearest running water; mean distance to nearest well (if present within 5m); presence/absence of leaf sponges. For each lobe, werecorded: length, wide, height (which multiplied gave

Despite intensive searching and monitoring over a
5-fold longer distance of riverbed and over a 6-fold longer
period in 2008, we found wells at a much lower rate andhad no direct observations of well digging. Thus, thestudy remained archaeological, that is, based on indirect
evidence.
Table 1 compares left vs. right lobes of bi-lobed tailings left at wells. As in 2006, there was no difference in2008 in volume between them. Similarly, the new variable, weight, also showed no difference between left
vs. right. Table 2 shows the extent of contrast between
volumes of left and right lobes, that is, the ratio of R/Lwhen right lobes were bigger vs. L/R when left lobes were
bigger. In both 2006 and 2008, there was no difference
between these ratios.
Table 1. Left vs. right ‘lobes’ of excavated sand tailings ofwells compared by volume for two years; weight of lobed
tailings only for 2008.

Year

N of
wells

2006
2008

25
54

Mean volume (cm3)
L lobe R lobe
10,679 11,759
7,030 5,854

p
NS
NS

Mean weight (g)L lobe R lobe p
NA
NA NA2,295 2,128 NS

L vs. R = Sign Test, two-tailed, NS = not significant; NA = not available

Table 2. Extent to which volumes of left vs. right tailings(lobes) (R/L, L/R) differ, for two years.

Figure 3. Measurement of water depth in wellcontaining leaf ‘sponge’.

Year

N of
wells

2006

25

2008

54

Dominant Lobe
Left
Right
Mean L/Rp
Mean R/L p
1.92
NS
2.22
NS
(n=31)(n=23)
2.00
NS
1.68
NS

L vs. R: Mann-Whitney U, two-tailed, NS = not significant

Pan Africa News, 20(1), June 2013
Table 3. Presence or absence of leafsponge tools at well, comparing two years.

Year
2006
2008

Tool
11
1

Well
No Tool80
120

Chi-squared test, two-tailed, p < 0.01

Total
91
121

7
Table 4. Relationship between rainfall (mm) and abundance of wells:
Month-by-month comparison of 2008 versus 1997-2006 (median).

Year(s)1997-2006
2008
Wells found 2008

Spearman’s rho, rs = -0.46, two-tailed, NS( )= extrapolated figure based on partial data only

However, there were differences between years: Table
1 shows that wells were smaller in 2008: Volume of bothleft and right lobes was less. Table 3 shows that in 2008,
only one leaf sponge tool was found at 121 wells, while in2006, one or more sponge tools were found on average atevery ninth well.

DISCUSSION

Rainfall (mm)
May June July Aug Sep Oct Nov Total
164
55
73
141 94 149 168 844(73) 98180
50
86 210 (182) 879
0
29
25
48
19
0
0
121

The symmetry between left and right lobes was replicated across study years, adding more data to the long
list of studies that have failed to find handedness in wild
chimpanzees (e.g. Marchant & McGrew 2013).
However, the unexpected inter-annual differences
found require explanation. Overall rainfall seems not to
suffice: Table 4 compares monthly rainfall totals over the
study period in 2008 with average totals for a previous 10yr period (1997–2006). Overall, 2008 was not an atypicalyear, although July was very wet (180 mm) and Augustvery dry (50 mm). There was no correlation between

monthly rainfall and abundance of wells.
Instead, the contrast may reflect hydrological differences, that is, the level of the water table in the riverbed.
A high water table allows for shallower wells and so,less sand needs to be removed, while a low water table
requires deeper wells and the removal of more sand.
Similarly, water can be drunk directly with the lips from ashallow well, while sponge use facilitates drinking from adeeper well.
Such hydrological factors rarely, if ever, have beenaddressed in field studies of wild chimpanzees, but they
may be crucial for explaining the presence or absence of
potable drinking water in hot, dry and open habitats, especially in the dry season. Further study is needed.

ACKNOWLEDGEMENTS

We thank: Uganda Wildlife Authority and Uganda Council
for Science and Technology for permission to do research at
Semliki; UWA rangers for field assistance and Semliki camp
staff for domestic assistance; Phineas Hodson and Samantha

Figure 4. Leaf ‘sponges’ for extracting water from wells. Top row: Intact leaves plucked from source. Middle row:
Processed leaves with softer tissue removed and used but stiff veins discarded. Bottom row: Used sponges.

Pan Africa News, 20(1), June 20138

adult female chimpanzee in the Bossou community.

Russak for research collaboration; Rebecca Jeanne Andrew
Memorial Fund, Miami University (THW) for funding for field
work; Leverhulme Trust (WCM) for funding for analysis andwriting up results.

Methods

Notes
1. Another population of chimpanzee (Pan troglodytes verus) in
southeastern Senegal has been reported to dig wells (GalatLuong & Galat 2000; Galat-Luong et al. 2009). The authors
reported that apes used tools to dig their wells, but baboons
as well as chimpanzees were said to dig wells, and no criteria
were presented for distinguishing between the two species’
wells. The first report is an abstract, and the second was
published in a non-peer-reviewed journal. Neither presented
any systematic or quantitative data, nor observations of
behaviour, so these claims must be taken as tentative until
documentation is provided.

Observation

REFERENCES

Galat-Luong A, Galat G 2000. Chimpanzees and baboons
drink filtrated water. Folia Primatol 71:258 (abstract).
Galat-Luong A, Galat G, Nizinski G 2009. Une consequence
du rechauffement climatique: les chimpanzes filtrent leureau de boisson. Geographia Technica. Numéro spécial,2009:199–204.
Hunt KD 2000. Initiation of a new chimpanzee study site
at Semliki-Toro Wildlife Reserve, Uganda. Pan Afr News
7:14–17.
Hunt KD, McGrew WC 2002. Chimpanzees in the dry habitats of Assirik, Senegal, and Semliki Wildlife Reserve,
Uganda. In: Boesch C, Hohmann G, Marchant LF (eds)
Behavioural Diversity in Chimpanzees and Bonobos.
Cambridge University Press, Cambridge, pp. 35–51.
Marchant LF, McGrew WC 2013. Handedness is more thanlaterality: lessons from chimpanzees. Ann NY Acad Sci
1288:1–8.
McG rew WC, Ma rcha nt LF, Hu nt K D 20 07. Et ho archaeology of manual laterality: well digging by wild
chimpanzees. Folia Primatol 78:240–244.

<NOTE>

Newly Observed Bird
Consumption by a Chimpanzee(Pan troglodytes) at Bossou,
Guinea, West Africa
Lira Yu1, 2, Boniface Zogbila3 &Tetsuro Matsuzawa1
1 Primate Research Institute, Kyoto University, Japan
2 Japan Society for the Promotion of Science
3 Institut de Recherche Environmentale de Bossou(E-mail: yu.lira.87z@st.kyoto-u.ac.jp)

Introduction

Bossou chimpanzees have shown low rates of meat
consumption compared to other communities (Sugiyama& Koman 1987; Hockings et al. 2012). Therefore, the addition of data describing their prey repertoire is imperative for ecological and behavioral studies at this site. Here,
we report a novel observation of bird consumption by one

Behavioral observations were conducted at Bossou,
one of the long-term research sites for chimpanzees located at the south-east corner of the Republic of Guinea,
West Africa (Matsuzawa et al. 2011). The core area usedby Bossou chimpanzees includes approximately 6 km 2,
dominated by primary and secondary forest and surrounded by cultivated or abandoned fields and shrub forest. Consumption of the wild bird was observed by LY
and BZ, who recorded all behavioral occurrences during
the event ad libitum.
On October 30, 2012, at 7:39 AM, we (LY and BZ)
observed four adult chimpanzees (JJ, TA, FF, and Jr) in
the forest. At 7:47 AM, three adult male chimpanzees (JJ,
TA, and FF) started to move. However an adult female
chimpanzee Jr (estimated to be 54 years old) remained.
For approximately ten minutes, Jr showed no movement.
At 7:57 AM, shrubs began to rustle. We heard a typicalcall of coucal (a bird belongs to cuckoo family) from anearby bush. The call consists of a series of ‘hoos’ which
at first increase in rate and volume and die away at theend (Serle et al. 1990). A few seconds later, Jr suddenly
moved toward the bush. We saw Jr capturing two nestlings at once from a nest that had been constructed in thebush and located ca. 2 meters above the ground. Jr walked
a few steps away holding the two nestlings one in eachhand and sat on the ground. Jr started to take a bite of thenestling in her left-hand. Then Jr put the remainder of thenestling on the ground and consumed a few leaves from anearby liana. We (BZ and TM) later identified the leaves
as Clerodendrum silvanum. This liana is characterized by
its needle-like spines that are derived from young stems
and petioles but not on epidermis (Hawthorne & Jongkind
2006). For the next few minutes, the nestling and leaves
were consumed in an alternating manner. Only the lefthand was used during consumption of both items, while
the right-hand continued to hold the other nestling. At
8:07 AM, Jr stopped eating and put what remained of theconsumed nestling on the ground. Jr stood up bipedally,
moved a few steps and started slapping the other nestlingon the ground. The nestling emitted some vocalizations
when it hit on the ground. In response, Jr released thenestling from her hand and stepped back. Soon, Jr took
the nestling again and continued to slap it on the ground.
After several cycles of slapping and stepping away fromthe nestling, Jr picked the nestling up and moved out ofsight.
At 9:09 AM, after we lost track of Jr, we returnedto the bush and discovered a bird carcass on the groundamid scattered leaves that had been consumed (see Figure1). We assumed this carcass was the first nestling that Jr
consumed because it lacked some body parts; its head andlegs were missing from the carcass while most of the bodytrunk remained intact. We could not find any detachedbody parts in the vicinity of the carcass, suggesting thatthese had all been consumed by Jr. Because of the emitted
call and the carcass, it was identified as the black-throatedPan Africa News, 20(1), June 2013

9

Figure 1. Discarded coucal nestling carcass on the ground. Head and legs were missing from the carcass.
(photograph by Boniface Zogbila)

coucal (Centropus leucogaster), a species of cuckoo family (Cuculidae). While cuckoos are known to be parasitic
in their nest habits, coucals construct their own nests andrear their young (Serle et al. 1990).

Discussion

This is the second confirmed species of bird preyed
upon by Bossou chimpanzees. Another species is thenocturnal West African wood-owl (Ciccava woodfordi)(Sugiyama 1989; Carvalho et al. 2010; Hockings et al.
2012). The prey reported here was a diurnal bird species,
the black-throated coucal. Fujimoto & Shimada (2008)
reported three cases of bird consumption at Mahale,
Tanzania, and reviewed the predation of wild birds inchimpanzees including other study sites; Bossou, Ndoki,
Kibale and Gombe. The current observation is in line withthese reports: The chimpanzee captured nestlings opportunistically upon detection and was solitary at the time.
After opportunistic prey-capture, chimpanzees arereported to display a variety of behaviors. At Mahale,
where predation on birds is observed more frequently
than at Bossou, chimpanzees show individual differenceswith regards to the captured birds: Some chimpanzees eat
them and others not (Fujimoto & Shimada 2008). In thisobservation, the chimpanzee at Bossou consumed the firstnestling immediately after capture, suggesting that the
chimpanzee may have recognized coucals as a prey species. Unfortunately, we could not determine whether or
not the second nestling was consumed.
The chimpanzee killed the second nestling more cau-tiously by slapping over time. Similar to the previously
observed predation of birds at Mahale, chimpanzees at
Bossou also sometimes discard captured animals without
consumption (Ohashi 2006), and sometimes is accompanied by toying (with the prey) behavior (Hirata et al. 2001;
Carvalho et al. 2010). Here, we saw the chimpanzee carrying the nestling after the slapping behavior. Hockingsand others (2012) noted that accompanying behaviors,
including grooming, plucking, rubbing, slapping, or carrying a captured animal, rarely occurs in adult chimpanzees at Bossou. Thus, the present observation added aninteresting case of predatory and accompanying behavior
by an adult female chimpanzee.

Acknowledgements

This research was financed by AS-HOPE of the Primate
Research Institute of Kyoto University and a JSPS Grantin-Aid for Scientif ic Research (#244525) to L. Yu and
MEXT #24000001 to T. Matsuzawa. We thank the Direction
Nationale de la Recherche Scientifique et Technologique(DNRST) and the Institut de Recherche Environmentale deBossou (IREB) for the collaboration and the permission to
conduct the research at Bossou. Special thanks are due to S.
A. Gaspard, the general director of IREB, and G. Yamakoshiof Kyoto University. Thanks are due to the local assistants,
especially H. Camara, for the identification of the bird andthe plant. Thanks are also due to M. Fujisawa who guided andhelped L. Yu during the stay at Bossou. We are also gratefulto M. Hayashi, Y. Yamanashi, M. Tomonaga, G. Ohashi, K.
Hockings, T. Humle, S. Carvalho, C. Sousa and A. MacIntosh
for helpful comments and edits on this manuscript.

10
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.
Fujimoto M, Shimada M 2008. Newly observed predation ofwild birds by M-group chimpanzees (Pan troglodytes schweinfurthii) at Mahale, Tanzania. Pan Afr News 15:23–26.
Hawthorne WD, Jongkind CCH 2006. Woody Plants ofWestern African Forests: a Guide to the Forest Trees,
Shrubs and Lianes from Senegal to Ghana. Royal Botanic
Gardens, Kew.
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.
Hockings KJ, Humle T, Carvalho S, Matsuzawa T 2012.
Chimpanzee interactions with nonhuman species in an anthropogenic habitat. Behaviour 149:299–324.
Matsuzawa T, Humle T, Sugiyama Y (eds.) 2011. The
Chimpanzees of Bossou and Nimba. Springer, Tokyo.
Ohashi G 2006. Behavioral repertoire of tool use in the wild
chimpanzees at Bossou. In: Cognitive Development in
Chimpanzees. Matsuzawa T, Tomonaga M, Tanaka M (eds),
Springer, Tokyo, pp. 439–451.
Serle W, Morel GJ, Hartwig W 1990. A Field Guide to the
Birds of West Africa. Collins, London.
Sugiyama Y, Koman J 1987. A preliminary list of chimpanzees’ alimentation at Bossou, Guinea. Primates
28:133–147.
Sugiyama Y 1989. Description of some characteristic behaviors
and discussion on their propagation process among chimpanzees of Bossou, Guinea. In: Behavioral Studies of Wild
Chimpanzees at Bossou, Guinea. Sugiyama Y (ed), Inuyama:
Kyoto University Primate Research Institute, pp. 43–47.

Pan Africa News, 20(1), June 2013

<NEWS>The Process of Creation of a New
Protected Area in the Democratic
Republic of Congo: The Case of
the Iyondji Community Bonobo
Reserve
Jef Dupain1, Andrew Fowler2, Phila
Kasalevo2, Tetsuya Sakamaki3,4,
Bongoli Lingomo5, Theo Way2,David Williams6, Takeshi Furuichi3,4,Charly Facheux2
1 African Wildlife Foundation, Nairobi, Kenya
2 African Wildlife Foundation, Kinshasa, DRC
3 Primate Research Institute, Kyoto University, Japan
4 Wamba Committee for Bonobo Research
5 Foret des Bonobos, Iyondji, DRC
6 African Wildlife Foundation, Washington DC, USA(E-mail: jdupain@awfafrica.org)

INTRODUCTION

Great ape populations across Africa are in crisis.
Their original habitat is decreasing rapidly, and only an
estimated 10% will be left undisturbed by 2030 if current
trends continue (Stiles et al. 2013). The rate of suitable
habitat loss for bonobos is particularly high, estimated at
29% between the 1990’s and 2000’s (Junker et al. 2012).
The total bonobo range area is about 418,803 km², but
only 97,975 km² is considered as suitable habitat (IUCN/SSC A.P.E.S. Portal http://apesportal.eva.mpg.de/). An estimated 42% of this suitable habitat is gazetted in protected areas (Campbell et al. 2012), with most of this within

Figure 1. Iyondji and other Maringa-Lopori-Wamba landscape protected areas.

Pan Africa News, 20(1), June 2013
the one Salonga National Park. In order to maximize theconservation of the cultural, genetic and ecological diversity of the bonobo, and to address the loss of critical
habitat, it is imperative to establish a network of protectedareas (PAs) that can sustain viable bonobo populations.
In 2012, as a next step towards such a network,
the African Wildlife Foundation helped gazette a new
bonobo protected area (PA), the about 1,100 km² Iyondji
Community Bonobo Reserve (Figure 1). The establishment of this bonobo PA and detailed information aboutbonobos living there were reported elsewhere (Sakamaki
et al. 2012). This article describes the significance of thisPA as a case study of the application of the Heartland
Conservation Process (HCP) (Henson et al. 2009) in one
macro zone in this landscape, and as a potential model forcollaboration between local communities, researchers anda conservation NGO.

I D E N T I F I C AT I O N O F T H E P R O P O S E D
PROTECTED AREA

The African Wildlife Foundation (AWF) has worked
in the 74,000 km² Maringa-Lopori-Wamba Landscape(MLW, CARPE, USAID) within the bonobo distribution
range since 2004 (Dupain et al. 2009). As a desired outcome in the MLW, AWF aims at the creation of a networkof interconnected protected areas covering 15% of thelandscape. To achieve this conservation goal, AWF utilizes an approach that was developed and implemented aspart of the overall Heartland Conservation Process. The
HCP balances science-based landscape scale conservation
planning with the sustainable aspirations and needs of
local people. Thus informed by a combination of habitat
suitability analysis, assessment of human threats and vulnerability to deforestation (Nackoney & Williams 2012),
consideration of future population-driven demand for land
(Nackoney & Williams 2013) and the outcomes of a defined public participation strategy, a conceptual model fora land use plan was developed (Dupain et al. 2008, 2010).
The plan created a sustainable vision for the landscapefeaturing various management zones including a networkof community conservation and PAs.
One PA zone targeted a forest block abutting the Luo
Scientific Reserve (Furuichi et al. 1998). Local communities were invited to a series of meetings as part of the public participation strategy for advancing the land use planning process (Sidle et al. 2012). These local communities
expressed their wish to realize this vision through theJapanese research team (Wamba Committee for Bonobo
Research-WCBR) and requested help from the WCBR
and AWF for the establishment of a bonobo community
reserve (Sakamaki et al. 2012).
Initial qualitative surveys were conducted and thefeasibility of the creation of a PA was evaluated (Likondo
et al. 2007; Likondo 2008). Evaluation focused bothon the biodiversity value and on the social context andthe attitude and expectations of the local communities.
Throughout the process, open consultation was organized with the local communities (the villages Yokali and
Yohala of the Groupement Iyondji). Land use and land
tenure was looked at in more detail (Kasalevo & Lingomo
2010). Subsequently, a Land Use Planning Strategy

11
Document was developed and implemented (Kasalevo &Dupain 2010). The research team from Kyoto University’s WCBR provided technical expertise for implementationof large mammal surveys with a focus on bonobos.

THE PROCESS OF GAZETTEMENT

Once the surveys confirmed the biological value
and the local communities had reached agreement on
objectives, AWF initiated and facilitated the the creationof the Iyondji Community Bonobo Reserve (ICBR) by
following the procedures as outlined in the Ministerial
Decree of April 2008: Décret n° 08/08 du 08 avril 2008
fixant la procédure de classement et de déclassement desforêts, Journal Officiel de la République Démocratiquedu Congo, 49ième année, nr special, 1ière partie 10 mai
2008, 7pp.
Local polit ico -ad m i n ist rat ive aut hor it ies(Groupement, Secteur, Territoire) consulted with the local communities and validated the draft management plan(see Kasalevo 2010). Following the government statutes,
a Provincial Consultative Council is expected to give its
opinion on management plans. However, in absence of
this council, the Provincial Governor gave his approval.
A document for creation of the PA was developed andpresented to the Ministry of the Environment and Nature
Conservation. This document featured results of biological and socio-economical surveys, the formal consent of
the communities, and the proposed geo-referenced limitsof the future PA, and was evaluated by a committee of experts assigned by the legal advisor of the Ministry. Upon
validation of this document by the aforementioned committee, the protected area was officially gazetted throughthe Ministerial Decree, signed on April 12th 2012 (Arrêté
Ministériel N° 020/CAB/MIN/ECN-T/15/ JEB/012 duportant classement de la Forêt de d’Iyondji).

DISCUSSION

The official gazettement of the ICBR in April 2012
brings the total area of bonobo habitat in formal protected areas within the 74,000 km² MLW landscape to
9,247 km² or 12.5%. With the Luo Scientific Reserveand Kokolopori Bonobo Reserve, ICBR contributes toa contiguous 5,627 m² bonobo conservation stronghold(see Figure 1). The process of creation of the ICBR is well
documented and represents the first PA created following the Ministerial Decree of 2008. The process proved
straightforward, feasible to replicate, and thus provides amodel for gazettement of other suitable PAs.
The identification and subsequent establishment of
the new PA in the MLW landscape combined sciencedriven spatial modeling with the aspirations expressedby local communities. The latter is only possible if thosecommunities are well informed on the processes to be
applied. For this, the public participation strategy was ofutmost importance. The transparent, collaborative effort
between local communities, researchers and conservation
NGO greatly improved the efficiency of this endeavor. For
example, having researchers on the ground allowed for
day to day monitoring of survey related activities, but alsoof the attitude of the local communities. For its part, AWF
had the conservation expertise and network to facilitate

12
the official procedures and contacts with local, provincial
and national authorities.
This new PA contributes to improved conservationof genetic diversity within the overall bonobo population(Kawamoto et al. 2013). The development of a network
of PAs throughout the bonobo suitable habitat will maximize the conservation of cultural, genetic, and ecologicaldiversity of the species. The ICBR, while not large in size,
advances these desired outcomes.
Going forward in the MLW landscape, AWF will focus on two directions:• Assure continuous connectivity between theanchor PAs to provide genetic linkages between
the region’s wildlife populations. Connectivity
zones will protect habitat between PAs from uncontrolled land use and land cover change. AWF,
in partnership with the University of Maryland,
continues its support of participative micro zoning in community managed forest between thecurrent PAs in the MLW landscape.
• Bring total surface of bonobo-habitat in PAs in
the landscape to 15%. At the time of writing, an
area has been identified through spatial modelingand consultations with the local communities
has commenced.
Additionally, it is critical to support the managementof the new PAs and assure their future sustainability/viability. The WCBR continues to provide technical support
for the habituation of bonobos at two field camps within
the PA (Sakamaki et al. 2012). AWF continues to support
the local NGO “Forêt des bonobos” and the DRC wildlife authority, the Congolese Wildlife Authority (ICCN)
in the development and implementation of a management plan for the Iyondji Community Bonobo Reserve.
Initiatives with other organizations and donors, e.g. IUCN
small grants program, are also taking shape. Throughthe process outlined herein, AWF hopes to mitigate thesteady loss of habitat and habitat suitability in the MLW
landscape and create a network of protected areas that cansustain bonobo populations.

ACKNOWLEDGEMENTS

This project would not have been possible without thepartnership with the Wamba Committee for Bonobo Research.
The same is true for the local NGO, Forêt des Bonobos, thelocal Iyondji communities of Yokala and Yofala, and theCentre de Recherche en Ecologie et Foresterie (CREF) of
the Democratic Republic of Congo. Support is given by the
Ministry of Environment and Nature Conservation and theMinistry of Scientific Research of the Democratic Republicof Congo. The present project was financially supported by
the US Fish and Wildlife Service Assistance Award (962000-G017 to AWF), Grants-in-Aid for Scientific Research of theJapan Society for the Promotion of 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-22-027, ITP-23-006
to T. Sakamaki), the Regina B. Frankenberg Foundation, andDisney Worldwide Conservation Fund.

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Warren Y, Williamson EA, Kuehl HS 2012. Recent decline
in suitable environmental conditions for African great
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Kasalevo P, Dupain J 2010. Iyondji Bonobo Community
Forest. Land use planning strategy document. AWF. Reportfor USAID/CARPE.
Kasalevo P, Lingomo B 2010. Draft of Report on the Studies
of Land Use and Land Patterns. AWF. Report for USAID/CARPE. http://carpe-infotool.umd.edu/IMT/LS9_MaringaLopori-Wamba/9010006_PA_Extention_Luo-Scientific_
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Kawamoto Y, Takemoto H, Higuchi S, Sakamaki T, Hart JA,
Hart TB, Tokuyama N, Reinartz GE, Guislain P, Dupain
J, Cobden AK, Mulavwa MN, Yangozene K, Darroze
S, Devos C, Furuichi T 2013. Genetic Structure of Wild
Bonobo Populations: Diversity of Mitochondrial DNA and
Geographical Distribution. PLoS ONE 8:e59660.
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de faisabilité d’extension de la Réserve Scientifique deLuo dans le bloc forestier du Groupement Iyondji, Secteurde Luo, Territoire de Djolu, District de la Tshuapa. AWF.

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United Nations Environment Programme, GRID-Arendal.
www.grida.no<BOOK INFO>Groups: The Evolution of Human
Sociality
Edited by Kaori Kawai
This volume is the product of a collaborative project based at the Research Institute for Languages andCultures of Asia and Africa at the Tokyo University ofForeign Studies. Researchers primarily involved in threefields—primate sociology and ecology, ecological anthropology and socio-cultural anthropology—came together
to discuss the shape and variations of groups as sympatricentities and the evolutionary historical foundations thathave led to the orientation of groups in present-day human society. To that end, the chapters in this volume turnto non-human primates for comparative purposes to consider the nature of the evolutionary historical foundationsof sociality.
In place of the past objective of “reconstructing” theecology and society of early humans, the works in thisbook instead aim to re-identify the creation and evolutionof that which is social and challenge the prevailing theoryof groups in socio-cultural anthropology. Specialists

13
on research into human beings and those studying nonhuman primates develop the debate about groups in thecontext of their own areas of expertise, at times in waysthat extend beyond the boundaries of their fields.

Hardcover: 426 pages
Publisher: Trans Pacific Press (April 1,
2013)
Language: English
Hardcover $109.500
ISBN-10: 1920901787
ISBN-13: 978-1920901783
Contents
Introduction—In Pursuit of an Evolutionary Foundationfor Human Society / Kaori Kawai
Part I: The Evolution of Sociality
1. The Sociology of Anti-Structure: Toward a Climax ofGroups / Kaoru Adachi
2. Assembly of Solitary Beings: Between Solitude and“Invisible” Groups / Motomitsu Uchibori
3. From Whence Comes Human Sociality? Recursive
Decision-making Processes in the Group Phenomenon
and Classification of Others through Representation /Kōji Kitamura
4. The Function and Evolutionary History of Primate
Groups: Focusing on Sex Differences in Locational
Dispersal / Naofumi Nakagawa
Article 1—A Group of Chimpanzees: The World
Viewed from Females’ Perspectives / Noriko Itoh
Part II: The Organization of Social Groups
5. The Ontology of Sociality: “Sharing” and Subsistence
Mechanisms / Keiichi Omura
6. Violence and the Autopoiesis of Groups: From theEthnography of Pirates and Feuds / Ikuya Tokoro
7. Forming a Gang: Raiding Among Pastoralists and the“Practice of Cooperativity” / Kaori Kawai
Article 2—Yesterday’s Friend is Today’s Enemy: The
Huli Society of Papua New Guinea / Masahiro
UmezakiPart III: The Formation and the Development of “We”
Consciousness
8. From the “Here and Now Group” to the “DistantGroup”: Hunter-gatherer Bands / Hideaki Terashima
9. Perceivable “Unity”: Between Visible “Group” and14Invisible “Category” / Toru Soga
10. The Small Village of “We, the Bemba”: The
Reference Phase that Connects the Daily Life Practicein a Residential Group to the Chiefdom / Yuko
Sugiyama
Article 3—The “Group” called the Kenya Luo: A Social
Anthropological Profile / Wakana ShiinoPart IV: Towards a New Theory of Groups
11. Collective Excitement and Primitive War: What is
the Equality Principle? / Suehisa Kuroda
12. Agency and Seduction: Against a Girardian Modelof Society / Masakazu Tanaka
13. Human Groups at the Zero-Level: An Exploration of
the Meaning, Field and Structure of Relations at theLevel of Group Extinction / Takeo Funabiki
Conclu sion — From “G roups” to “I nst it ut ions”:
Summary and Prospects
Epilogue—The Legacy of Hitoshi Imamura: The Macrolies in the Micro / Ryōko Nishii
Notes
Bibliography
Index<FROM THE EDITORS>
It is our pleasure to reach the 20th year of Pan Africa
News (PAN), which was founded in the summer of 1994.
We would like to express our sincere gratitude to everyone who has generously supported us as readers, contributors, and reviewers.
The late Professor Toshisada Nishida, the first Chief
Editor, launched PAN in response to an idea generated at the conference “Understanding Chimpanzees”,
which took place in Chicago, in November 1986. Dr.
Jane Goodall and other chimpanzee experts felt that theyneeded a newsletter in which fieldworkers studying wild
chimpanzees at various sites could share their information and experiences. At that time, the Gombe and Mahale
projects had accumulated research experience spanningmore than two decades, and many other projects exploring the two Pan species had emerged across Equatorial
Africa. However, researchers were not always familiar
with on-going studies and the conservation status ofchimpanzees at sites other than their own. All they could
get was relatively limited information through papers
published in certain journals with wider readerships. PAN
was intended to serve as a forum where fieldworkers interested in the Pan species could communicate informallyto facilitate their research and conservation efforts.
For the past two decades, PAN has successfully lived
up to expectations, but its role seems to have expanded.
In 1997, we adopted a new editorial system for managing
the submission and peer-review process to meet the needsof fieldworkers wanting to publish small findings about
the Pan species, given that other scientific journals, evenprimatology journals, tend not to publish papers based on

Pan Africa News, 20(1), June 2013a single or rare observation. Subsequently, the diversity ofwild chimpanzee behaviours in different areas became a
hot topic, interesting many researchers. As a result, many
good papers with invaluable findings, raising important
issues, have appeared in PAN and, due to our open-accesspolicy, have circulated within the scientific communityand been cited frequently in other journals.
As we enter a new decade of PAN, we feel the needto change several of its elements so that we can maximize
the niche that this journal has occupied. Such changes andimprovements could include the guidelines for authors,
editorial policy, publishing schedule, page layout, andjournal title/description.
We might also widen the scope of this journal.
Although we will keep our old policies regarding publishing information confined to chimpanzees and bonobos in
the wild or in sanctuaries in Africa, it might benefit PAN
readers if we publish papers on other species and news
that is relevant to research and conservation of the Pan
species. Next, we might start online publication on the
day of acceptance, because the current biannual publication can keep early contributors waiting for 5~6 months
until they see their accepted papers. Third, we should
define policies for research ethics that we should observe
while studying wildlife in Africa. We want to make such
improvements in time for the next issue. Any suggestions
will be appreciated.
We will post an update of each change online, once
it has been fixed. Please visit our website (http://mahale.
main.jp/PAN/) before you submit your manuscript.
Some editing changes have been applied to this issue.
We have changed the citation style. We have abandoned
the original style of superscript numerical citations in the
text and adopted the more common style of the author’s
surname and publication year. References at the end of
the paper should be listed alphabetically by the author’s
surname.
From the next issue forward, the deadline for manuscripts will be 1 month earlier; it has changed from theend of November to the end of October, which enables us
to ask authors to correct their proofs. When you submit
your manuscript, you can suggest two potential expert
reviewers. You can also suggest researchers to whom
we should avoid sending your manuscript for reasonable reasons. For further details, please see the online
information.

Pan Africa News, Vol. 20, No.1
Published in June, 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)