Anda di halaman 1dari 464

WORLD ATLAS OF

(iKEATAPtS
AND THEIR CONSERVATION

'

%
\

ROYALTIES FROM THE SALE OF THIS BOOK WILL SUPPORT THE CONSERVATION EFFORTS OF THE GREAT APES SURVIVAL PROJECT

EDiTEDBY

JULIAN CALDECOTT

and

LERA MILES

foreword by

kofi a.

annan

fAt
t^ms
^Oj
UNEP

WCMC

ESPITE THE DEDICATED EFFORTS OF

MANY INDIVIDUALS

and organizations, the great apes our closest


tives-are

in

danger

of extinction. This sw/eeping atlas provides

a comprehensive overview of what

all

six

species of great

is

currently

Created

Apes

in

known about

apeschimpanzee, bonobo, Sumatran

orangutan, Bornean orangutan, eastern


gorilla.

living rela-

gorilla,

and western

association with the United Nations Great

Survival Project

(GRASP),

this

background on behavior and ecology

book gives a thorough

for

each species,

includ-

ing detailed habitat requirements, the apes' ecological role,

and the possible consequences


of Great

Apes

also offers a

rent conservation efforts,

each species across


illustrations

make

the

its

full

of their decline.

World Atlas

description of the threats, cur-

and additional protection needed

entire range.

abundance

for

Many full-color maps and

of information accessible to

a broad readership, from specialists and policymakers to general

readers concerned about the survival of these charismatic

primates.

This book represents the work of a dynamic alliance of

many

of the world's leading great

ape research and conser-

vation organizations. Bringing together United Nations agen-

cies,

governments, foundations, and private-sector interests,

the project aims to raise the international profile of great ape

conservation and to build the

Readers
tions

in

vation

learn

what work

is

political will for further action.

being done by specific organiza-

support of great ape conservation and where conser-

is

most needed and most

likely

to

be

effective.

World Atlas

of

GREAT APES
AND THEIR CONSERVATION

'

^f^^j^'^*^
^^,i.M3m
jsI^^I^ShHR^Bh
-

MM^BKrirarr'^^

i^mm^."'S^-i

'

'^^^'^^.>^^^^^^^r

^-S^^^

-^-"-^

'vJ^osHh

^ppi^^/

L-

'

-'>^f^';S..,-'v'-*^

%'^^^p^^^!^,

.^

'

^^J'.^'^Pholograph by Sebastiao Salgado/Amazonas images

Published

in

UNEP-WCMC

association with

by the University

of California Press

University of California Press

Berkeley and Los Angeles, California


University of California Press, Ltd.

London, England

2005

UNEP World

Conservation

Monitoring Centre

UNEP-WCf^C
219 Huntingdon Road

UK

Cambridge. CB3 0DL.

+a 101 1223 2773U


Fax: +W 101 1223 277136

Tel:

E-mail: info@unep-wcmc.org

Website: www.unep-wcmc.org

No

part of

ttiis

book may be reproduced by any means, or

transmitted or translated into a machine language witfiout


the written permission of the publisher

The contents

of this

or policies of

UNEP-WCMC.

volume do not necessarily

views

reflect the

contributory organizations, editors, or

publishers. The designations employed and the presentations do


not imply the expression of any opinion whatsoever on the part of

UNEP-WCIvIC or contributory organizations,

editors, or publishers

concerning the legal status of any country, territory,


or

its

authority, or concerning the delimitation of

boundaries, or the designation of

its

name

its

city,

or area

frontiers or

or allegiances.

Clothbound edition ISBN: 0-520-24633-0

Cataloging-in-Publication data

is

on

file

with

the Library of Congress.

Citation: Caldecott,

and

J.,

Miles,

their Conservation.

L.,

eds 120051 World Atlas of Great Apes

Prepared

at the

UNEP

World Conservation

Monitoring Centre. University of California Press, Berkeley, USA.

World Atlas

of

GREAT APES
AND THEIR CONSERVATION
.,

'

40-E

20'E

O'E

60*E

lOO'E

80-E

..

20"N-

20-N

0"N

-20-S

ill'-

Great ape distribution

^M

Bonobo

20'S1

Chimpanzee

^|

0"E

20-E

Gorilla

Orangutan

000

BO'E

60'E

41D*E

4 000

6 00 Okm

lOO'E

EDITED BY

^^

120"E

JULIAN

CALDECOH

and

Foreword by

UNIVERSITY OF CALIFORNIA PRESS


Berkeley Los Angeles London

120'E
^

LERA MILES

Kofi A.

Annan

^)
'W

UNEP WCMC

m
-

World Atlas

of Great Apes and their Conservation

World Atlas

Great Apes

of

and their Conservation


SUPPORTING ORGANIZATIONS

Prepared at

UNEP

World Conservation

Monitoring Centre

The United

219 Huntingdon Road

Programme

Vk^'^^

Cambridge CB3 DDL. UK

body

Nations

Website; www.unep-wcmc.org

the principal United

the

in

environment.
leading

Environment

Nations
is

role

Its

global

field

of

to

Is

the

be the

environmental au-

sets the global environmental agenda,

that

Editors

thority

Julian Caldecott

that

Lera Miles

environmental dimension

promotes the coherent implementation


of sustainable

of

the

development

within the United Nations system, and that serves as an

Cartography

authoritative advocate for the global environment.

Lee Shan Khee

objectives include analysis o* the state of the global

Matthew Doughty

environment and assessment

Mary Edwards

environmental trends, provision

of global
of

Its

and regional

policy advice

and

early warning information on environmental threats,

and promote international cooperation

Research assistant

and

Brigid Barry

and action, based on the best

to catalyze

scientific

and technical

capabilities available. Website: wAvw.unep.org

Production editors

Helen de Mattes

Angela Jameson

^B

QclrQ

Laura Kirby

for

Tim Osmond

environment

quality of

life

at

home and

internationally,

thriving

economies and communities

in

rural areas

and

a countryside for all to enjoy. Website: www.defra.gov.uk

The Ernest Kleinwort Charitable Trust


Origination

Swaingrove Imaging

banson@ourplanet.com

better

and sus-

economic prosperity

Raul Lopez Cabello

17e Sturton Street

now and

other industries that meet consumers' requirements;

Layout

Cambridge CBl 2QG, UK

for

better

through sustainable farming, fishing, food, water, and

Dormon

A Sanson Production

working

for everyone,

tainable use of natural resources;

Index

Printed and bound by Butler and Tanner,

Environment,
is

generations to come. This includes a

Valerie Neal

Jill

for

Food and Rural Affairs

sustainable development: a

'~'""-""-''

Jane Lyons

The UK Department

UK

World Atlas

of Great Apes and their Conservation

Acknowledgments

would

editors

Themense

the resources needed to

we must thank

reality. First,

lent

their

record their im-

to

Like

who committed
make this atlas a
organizations who

gratitude to all those

the

financial support:

Environment Programme Division


Conventions lUNEP DECI and

Nations

United

the

Environmental

of

Division

Early

of

Warning and Assessment lUNEP DEWAl; the UK

Department

for

and the Ernest Kleinwort Charitable

(Defral;

and other data are named

we thank them

chapters, and

in

in this

their creators,

of

the individual

again here. Their help

has made a tremendous difference. Most

images

Trust,

peer reviewers, and providers

Authors,
spatial

Environment. Food and Rural Affairs

of the

mental

in

getting the project started.

and Lucy Fish provided much support

May

cartographers. Ian

instru-

Simon

Blyth

to

set up the interactive

the

map

service that helped reviewers to audit the data.


Pragati Tuladar helped to locate

named

in

some

of the places

Simon Burr and Maria Murphy

the text.

helped us with the logistics of the peer review. Mary

Cordiner helped us to obtain various


literature. Brigid

vital

pieces of

Barry and Lee Shan Khee each

devoted months to the book, Brigid concentrating

on the

text

and photos, and Shan Khee on the maps.

our thanks

Jerry Harrison,

are credited alongside each one.

David Jay, Tim Johnson, Rebecca Kormos, Mark

may

Leighton, Kirsty Mackay, Daniel Malonza, Corinna

in

in

diverse

the form of

ways

names

that

associated

networking, providing introductions and


inputs, by helping with

mundane

but

essential tasks, or by providing moral support at


critical times.

Virtually

everyone

and the GRASP Secretariat


in

in

at

UNEP-WCMC

Ravilious,

Ian

to Phillip Fox,

Redmond, Melanie

Woods, and Kaveh Zahedi

We hope
generosity of

equal interest

new and

that this

all involved,
in

for their

book does
and

Virtue,

Matt

ongoing support.

that

it

justice to the
will kindle

great ape conservation

an

among

larger audience.

Nairobi, as well as

the nongovernmental conservation

organizations within the

lUCN/SSC Primate
Great Apes,

Kim McConkey, and Adrian Newton were

who

with particular sections - through strategic conver-

many people

thanks as follows.

offer particular

to

Finally,

Many others contributed

anonymous

like

Jared Bakusa, Brian Groombndge, Florence Jean,

book were generously contributed by

not be reflected

sations,

would

GRASP network and

Specialist

the

Group Section on

falls into this category.

With deep apologies for any omissions,

we

Julian Caldecott

and Lera Mites

Digitized by the Internet Archive


in

2010 with funding from

UNEP-WCMC, Cambridge

http://www.archive.org/details/worldatlasofgrea05cald

World Atlas

of Great Apes and their Conservation

Foreword
Kofi A.

great apes are our kin. Like us, they are

The

self-aware and have cultures, tools,

and medicines. They can learn

politics,

use sign

to

language, and have conversations with people and


with each other Sadly, however,

them

with

the

we

have not treated

respect they deserve, and their

numbers are now

declining, the victims of logging,

disease, loss of habitat, capture, and hunting.

Nevertheless there are signs

some

places,

governments have taken the lead

conservation efforts, often cooperating


national frontiers.
that

whoever

It

has been achieved and what we must do

apes better when they


result of education,

actions,

be

treat

each other

better,

as a

good governance, and reduced

poverty But saving the great apes

is

also about

we can
many people who
water, and much else.

also protect the livelihoods of the

on forests for food, clean

In

Indeed, the fate of the great apes has both practical

in

and symbolic implications

across

central

it

the

saving people. By conserving the great apes,

beings to

move

human

for the ability of

to a sustainable future.

Great apes cannot be conserved for free. The

has become increasingly clear

initiates

if

great apes are to survive. Often, people treat great

rely

hope.

of

Annan

Great Apes Survival Project documented

in

this

governments, local governments, international

publication can help by mobilizing resources. But

nongovernmental

this

citizens, local

they

who

need

to

live

organizations,

communities need

to

or

individual

be involved.

with the great apes, and

it

is

they

It

who

have the incentives - such as sharing

revenues from tourism This

atlas

conservation.

It

tells

to

the

is

in

of

great

only part of the answer, and other good ideas


to protect the great

We need

ordinary people

and protect them.

companies

conserve them.
story

is

on how

ape

describes both the progress that

they

live.

We

to 'adopt'

We

need

in

apes are also needed.


their millions to love

need governments and

them and

the places

where

to turn the tide of extinction that

threatens our nearest

living relatives.

Kofi A.

Annan

Secretary-General of the United Nations

World Atlas

of Great Apes and their Conservation

Contents
Acknowledgments

Gorilla overview

Western

FOREWORD

Map

Annan

Kofi A.

GonY(a9on7(a|

Western lowland

7.1

maps

The Cross River

7.1

10

[Gorilla gorilla diehli]

12

distribution

Map: Cross River

INTRODUCING GREAT APES


Richard Leakey

Box

Evolution, dispersal, and discovery of

Box

13

the great apes

makes

1.1

Box

1.2 Cryptic

Wtiat

world

105
106

a primate?

Box

18

apes

109

109

gorilla

window

113

of gorillas

7.3 Potential medicinal value

of gorilla

Box

gorilla

7.2 Forest clearings: a

into the
1

gorilla

distribution

Box
Using the

gorilla

97

1 1

foods

7.4 Gorilla

censuses

124

26
8

GREAT APE BIOLOGY

Eastern gorilla (Con'//a ber/nge/l

Map

29

Jane Goodall

Box

Eastern gorilla distribution

8.1

8.1

Coexistence of gorillas

129
130

137

and chimpanzees
2

Great ape habitats: tropical moist

31

forests of the Old World

Map

2.1

Great ape tiabitats

of

Sumatra

2.2 Great

liabitats of Africa

Chimpanzee and bonobo overview


Box

Use

3.1

of

8.3

human languages

in

gorillas

The vocal behavior

of

138
142

gorillas

Box 8.4 Eastern


ape

gorilla

tourism

150

36
9

8.2 Infanticide

Box

mountain

32

and Borneo

Map

Box

Orangutan overview

153

43
by

46

10 Bornean orangutan (Pongopygmaeus)

Map

captive great apes

10.1

Bornean orangutan

161

162

distribution

Chimpanzee Pan frog/odytesl


4.1

Chimpanzee

Box

53

Map

54

distribution

Box

4.1

Chimpanzees as predators

58

Box

4.2

Chimpanzee

64

Box

4.3

Chimpanzee cultures

66

Box

4.4

Seed dispersal by chimpanzees

70

Box

4.5 Reintroduction of

vision

orphan

to

Box
Box

76

83

5.1
5.1

Bonobo

distribution

Seed dispersal by bonobos

and the survival


Box

5.2

in

degraded

169

0.3 Peatlands in

Southeast Asia

76

Sumatran orangutan (Pongoa/)e<//l

Map

11.1

Sumatran orangutan

185

186

distribution

Bonobo Pan pan/'scus)


Box

Orangutans

as habitat for orangutans

Box

Map

10.2

165

dipterocarp forests

habitats

11
for

tourism

Adaptations of bearded pigs

life in

74

chimpanzees
Box 4.6 Chimpanzee habituation

10.1

of

ram

84
87

11.1 Culture

and

sociality in

192

Sumatran orangutans
Box 11.2 History

of the

Leuser

200

Ecosystem

forest

Bonobo communication

90

12 Gibbons: the small apes

205

World Atlas

CONSERVING THE GREAT APES

215

Map
Map

13 Challenges to great ape survival

Box 13.2

217

Lucky gorillas?

221

Satellite analysis of threats

223

13.1

227

Bushmeat hunting and trade

228

Box 13.5 Ebola and great apes

in

231

234

13.1 Historical conflicts

affecting the African great

Republic of Ghana

Conservation measures

in

play

Orangutan tourism

14.1

Chimpanzee

16.10

Chimpanzee

16.11

Republic

Chimpanzee

Box 14.2 Sendje, an orphaned

242

264
268

Map
Republic

distribution

Chimpanzee

distribution

Box

15.1

Great apes, consumers,

276
280

Map

Rwanda

Republic

WHERE ARE THE GREAT APES AND


WHOSE JOB IS IT TO SAVE THEM?
Ian

286

Chimpanzee

Map

16.18
of

Chimpanzee

Chimpanzee

Map

distribution

293

Republic of Angola

293

Map

16.1 Great

ape distribution

Republic of Burundi

Map

16.2

Chimpanzee

distribution

Cameroon
Map 16.3a Chimpanzee distribution
Map 16.3b Gorilla distribution

Republic of

16.20 Great ape distribution

Map

16.4 Great

ape distribution

Republic

Map

16.5 Great
of

ape distribution

16.6

Chimpanzee
of

Republic of Indonesia

300

305
306

Map

401

406

412
distribution

17.1

Orangutan distribution

17.2

Orangutan distribution

414

417
418
425

Malaysia

Map

396

417

17 Asia

299

426

308

AFTERWORD

431

Russell A. Mittermeier

316

the

322
328

Cote d'ivoire

Democratic Republic

Chimpanzee

392

320

Republic of the Congo

Map

16.21

294

314

Central African Republic

Map

384

404

United Republic of Tanzania

16 Africa

379

400

Republic of Uganda

Redmond

372

395
distribution

Sudan

16.19

367

391
distribution

Republic of Sierra Leone

Map

and the media

363

383

16.16 Great ape distribution

16.17

358

378

Republic of Senegal

15 Lessons learned and the path ahead

354

371

16.15 Great ape distribution


of

350

366

Chimpanzee

Federal Republic of Nigeria

Map

chimpanzee

16.14

343

362
distribution

of Liberia

16.13

342

357
distribution

Republic of Guinea-Bissau
16.12

336

353
distribution

Republic of Guinea

Map
Box

334

348

16 9 Great ape distribution

Republic of Mali

range states

ape distribution

Republic of Gabon

Map

ape

Equatorial Guinea

16.8 Great

Map

Central Africa

Map

Map

Map

Sendje, Equatorial Guinea

in

16.7b Gorilla distribution


of

Map

traditions
13.4

Republic

Map

Gombe chimpanzees
Box 13.3 Human beliefs and
to

Box

Bonobo and chimpanzee

16.7a

distribution

Toshisada Nishida

Box

of Great Apes and their Conservation

distribution

329

Congo

332

Annex: Great Apes Survival Project

GRASP and
Index

433

Partners

439

World Atlas

of Great Apes and their Conservation

maps

Using the
maps

The

in

book show forest cover,

this

observations

In

was

the Salonga National Park, because

designated areas, and species distributions.

Much

consult the species chapter or country profile text

information

is

condensed

into

each

map, so some explanatory notes are given here.

The

possible range of the species or subspecies.

'confirmed ranges' are smaller areas within which


is

known

be present. The point data

to

show species presence (squares


presence

I?),

exception

known

or

of

or circles], alleged

local extinctions

these

extinctions,

observations rather than

With the

(X|.

maps

are

of definitive

species

maximum

shown. The estimated ranges' are the

carried out there

for information

For Africa, three types of species data are

the species

survey

of

presence or

in

about the relative density

is

based on

natural forest. Park and reserve data are

The dates shown

in

date of last observation.

there are no

because the

recent

site

the legends indicate the


If

the date

an old one,

is

may

but this

records,

has not recently been

be

highest

level

designation

of

official

the species.

Where

a species has definitely

from a location,

lost

this

shown

is

been

explicitly.

protection

la

associated

have

some form

an

with

UNESCO

shown

national
to IV or a

International

World Heritage

Ramsar

of designation not

Sitel;

covered by the

Abbreviations have been used for the designation

names, such as FR

for Forest Reserve.

These are

listed opposite.

The African maps present

a compilation of

recorded observations and estimated range areas


of

great apes, put together by

Thomas

ButynskI

(Conservation International! and updated at

WCMC

have occurred since 1940.

searchers worldwide. The Southeast Asian

unless

all

that

species

is

It

The density

of

under-represented.

observation points

to Indicate the actual density of

as

likely to

example.

say

been destroyed, so

suitable habitat has

local extinctions are inevitably

to

difficult

no longer present

is

apes;

Is

it

is

Map

5.1

shows

a large

number

Illustrate the forest blocks in

together with their estimated density.

present,

ing the Population

Workshop

recent

maps

which orangutans are

These data were put together by researchers attend-

at least

of

UNEP-

with help from conservationists and re-

unlikely

indicate density of survey effort. For

Site

those that

Extinctions include only those that are recorded to

definitively

held

In

2004, and by the


Unit

in

and Habitat

Viability

Assessment

Jakarta, Indonesia,

team

at the

In

January

Leuser Management

Sumatra, Indonesia (see Chapter

111.

KEY TO ALL BACKGROUND DATA ON MAPS


Protected areas (boundary unknown)

Nationatly (lUCN Cat, l-IV) or internationally protected area

Other designated area

Proposed as

Other features
National capital

a protected area

Other

Tree cover (percent)

CHo
i-io

city

International boundary

Primary or secondary road

11-z.n

41-60

61-100

Protected areas [boundary delineated!


River
I

Nationally IIUCN Cat.

Other designated area

l-IVl

or mternationatly protected area

Proposed as a protected area

10

in

have the

above; and those that are proposed for designation.

visited by a

researcher rather than because of the absence of

to

designation assigned to lUCN Category

or Biosphere Reserve, or a

include those of great ape nests, tracks, or

known

three categories: those areas

within the estimated ranges shown.

mapped

of

plantations and degraded forests as well as Intact

convention, such as a

other signs as well as of the animals themselves.

shades

in

imagery, and includes

satellite

absence: presence should not be ruled out anywhere

The observations

of the

different parts of the range.

The tree cover shown on the maps


green

Please

2004.

in

Water body

World Atlas

ABBREVIATIONS USED ON THE MAPS


Biosphere Reserve
BR
Conservation Area
CA
Classified Forest
CF
Commercial Forest Reserve
CFR

NP
NR

National Park

NHM

National Historic

PF
PFR
PL

Protection Forest

Rehabilitation Center

Nature Reserve

Monument

FaR

Faunal Rese^^(e

FFR

Forest and Floral Reserve

FNR

Forest Nature Reserve

FR

Forest Reserve

RC
RR

GR
HA
HP
HR

Game

RS

Ramsar

Site

Reserve

for Scientific

Hunting Park

RSR
ScR

Scientific

Hunting Reserve

SNR

Strict

HZ

Hunting Zone

SR

Special Reserve

lER

Integral Ecological Reserve

WHS

World Heritage

NCU

Nature Conservation Unit

NF
Non-HFR

National Forest

WMA
WR

Non-Hunling Forest Reserve

WS

Reserve

Hunting Area

of Great Apes and their Conservation

Protection Forest Reserve

Protected Landscape

Resource Reserve
Research

Reserve

Nature Reserve

Wildlife

Site

Management Area

Wildlife

Reserve

Wildlife

Sanctuary

MAP DATA SOURCES


Protected areas

lUCN

119941 Guidelines for Protected Area

Management

Categories. lUCN, Cambridge,

UK and

Gland, Switzerland.

http://vvww.unep-wcmc.org/protected_area5/categor1es/eng/index.html.

World Commission on Protected Areas 120041 World Database on Protected Areas. UNEP-WCMC. http://sea.unepwcmc.org/wdbpa/index.htm. Accessed September 2004.

Tree cover
Hansen, M., DeFries,

R.,

Townshend,

J.R., Carroll, M., Dimiceli,

Continuous Fields. Global Land Cover


vcf/.

Facility,

C, Sohlberg,

R. (20031

500m M0DI5

Vegetation

College Park, Maryland, http://glcf.umiacs.umd.edu/data/modis/

Accessed September 13 2004.

Rivers
Petroconsultants (CES) Ltd (1990) Mundocart/CD: Version

2.0.

Petroconsultants ICESI Ltd. London.

Roads, country boundaries, coasts, inland water bodies

DMA

(1992) Digital Ctiart of the World. Defense Mapping Agency, Fairfax, Virginia.

Cities

ESRI (20031 ESRl Data

See the

map

data sources

Citations are
lists

& Maps

publications/WAGAC. A

maps

the country profiles

numbered separately

are not included

for the

in

2003. ESRI. Redlands, California.

in
list

this

in

in

Chapters 16 and 17 for great apes data sources.

each chapter and country

profile.

volume, but can be accessed online

of further

reading

is

The corresponding numbered reference

at;

http://vvww.unep-wcmc.org/resources/

given at the end of each chapter and country profile. Data sources

are also listed and usually represent additional further reading.

11

World Atus

of Great Apes and their Conservation

Introducing
great apes
Richard Leakey

became personally aware

threatened as they are today.

Mary Leakey discovered


remote

fossil

Victoria.

The

to

from

As an

my

my mother

africanus,

in

had

in

this

name Proconsu/ was

And so

chimpanzee

captive

was

it

that

chimpanzees as well as

apes became imprinted on

Much

my

later,

was

my young

fossil

who was

very involved

getting Jane Goodall established on her wild

chimpanzee studies
Republic
city

of

that

Gombe

at

Tanzania. Jane's

in

United

the

work and the

was generated through

publi-

National

the

of

in fact

one

is

talk of

seven. That six

This atlas of great apes

is

to

being the

timely and sets out

a great deal of information that

ignorant

of.

The threats

to

bonobos, and chimpanzees are


of habitat

many people

are

orangutans, gorillas,

many

but the loss

as remaining forests are plundered

surely a major concern. Disease too

is

is

a worry,

where there are increasing contacts

through tourism.
I

believe

we have

descendants as well as

an
to

obligation

to

our

our ancestors: the

remaining wild great ape populations must be


protected for

all

time.

As humans, we need

advance a new moral imperative

to

survival of these wonderful relatives,

more than anything

know them

to

We

which are

a sad testimony

Geographic Society and other media probably did


else to alert the general public

obvious

intelligent of the set.

particularly

brain.

further 'ape conditioned' by

father, Louis Leakey,

mercy

most

London Zoo.

the

at the

and a poor reflection upon our claim

inspired by Consul,
in

should be

all of

the

was about and

is

something separate.

ape species remaining,

day

was

sifying ourselves as

threatened, while there are

in

it

fundamental error was made when clas-

Lake

field

old,

that a

The great

hairy abstract beasts.

six great

learned about apes, especially fossil

famous

year

five

the fuss

all

some

not simply

apes and ourselves are so close that

then

be flown

to

mother's arrival

inquisitive

know what

to

ones! The

in

9i9

England for study, and the press had a

anxious

Rusinga Island

on

site

In

7 million year old find

covering the skull and


country.

was

a fossil skull of a primitive

known as Proconsul

ape,

apes when

of

and long before they were as

very young

better

is

to

ensure the

and getting

to

surely a good start.

the existence of chimpanzees and, by extension,

the other great apes.

time, other people and

In

other studies continued to enthral the public and so


build an

awareness

of

our closest relatives

living in

the shrinking forests of Africa and Southeast Asia.

Molecular biologists and geneticists have

how

demonstrated
to

chimpanzees

orangutans.

measures
and

to

gorillas

we

are

and

that in calling for

remaining populations

improve on the care and husbandry

in captivity,

12

protect the

we

close

bonobos,

must be stressed

It

to

biologically

and

are speaking of our

own

of

those

relatives.

Richard Leakey

Evolution, dispersal, and discovery of the great apes

Chapter

Evolution, dispersal, and

discovery of the great apes


Martin Jenkins

family Hominldae

The

one

is

mammals,

families of

smaller

of the

with seven

living

species. Six species are confined to various

wooded

forested or
II.

e.

habitats

In

the Old World tropics

the tropical parts of Africa and Euraslal;

beringei] and the western gorilla [Gorilla gorilla],

while

orangutans are separated

the

Sumatran

[Pongo

orangutan

abelii]

There

all of

is

second group

of living

gibbons, which are briefly covered

some critically so. The seventh

These are generally placed

Is

ubiquitous

and enormously abundant, probably the most

Homo

large animal that has ever lived:

numerous

the
the

Bornean orangutan [Pongo pygmaeus].

these are considered under threat of extinction,


species

into

and

the Hylobatidae, although

them

too as

members

in

apes; the

Chapter

some taxonomists

of

12.

a separate family,

in

regard

the family Hominldae.

species

four genera

are

sapiens, our

Currently about

activities of

recognized, ranging through Southeast Asia from

Is

own species. It Is entirely thanks to the


humans that each of the other species

currently

such a precarious

In

now

however,

We

state.

the unique position where.

in

have the collective

we can

will,

are,

we

If

reverse this trend

and ensure that our closest relatives have a viable


future on the planet. This

which

a course by

Apart from

this

volume attempts

to chart

beings [Homo] there are

three genera of great apes alive today; gorillas

chimpanzees

[Goriita],

The

[Pongo].

first

while the third


recently, there

two

of

[Pan],

occurs

Southeast Asia. Until

in

were generally accepted


gorilla],

[the robust or

to

be one

two species

common

of

chimpanzee.

Pan troglodytes and the pygmy chimpanzee or

Pan paniscus],

bonobo.

and

one

species

of

orangutan [Pongo pygmaeus]. Recent studies and

changes

In

approach

to

taxonomy have

led to the

populations of gorilla and orangutan each being


classified

as two separate species.^'

It

is

this

taxonomy, as endorsed by the Primate Specialist

Group
Is

of

used

lUCN-The World Conservation Union,


In

this

book, although

universally accepted.""

considered

to

it

Hence the

comprise the eastern

a lineage with the other

the forests of the orangutans, but

unique features

of lifetime

territoriality within

their

hands

like a

show

monogamy, duet

small

home

movement

own

singing,

ranges, and a
by swinging from

pendulum beneath

Many gibbon species

apes and

their

tree branches.

are also highly endangered.

orangutans

these are confined to Africa,

species of gorilla [Gorilla

chimpanzee

and

in

south China, Borneo, and Sumatra.

to Java,

The gibbons share

specialization for rapid

might be achieved.

human

Assam

12

that

has not been


gorillas

are

gorilla [Gorilla

DISCOVERY OF THE GREAT APES

long history

Apes and humans are no strangers


Cave deposits

in Viet

half a million years

Nam

erectus,'^

each other.

around

ago Imyal contain the remains

orangutans mingled with those

Homo

to

that date from

while cooked

of

the early

of

human

orangutan bones

dating to around 35 000 years ago have been found


in

the Great Caves of Niah

Africa,

although direct

in

fossil

Sarawak, Borneo.
evidence

Is

In

lacking,

humans and apes have undoubtedly shared the


same forests for millennia. To the western world,
however, these creatures remained half known
and

little

Indeed

it

understood

was

until

comparatively recently.

not until the early 20th century that

the last species

was described

scientifically

and a

13

World Atlas

of Great Apes and their Conservation

reasonably clear picture of this previously enigmatic

group

animals emerged.

of

The word ape' was applied


a

number

macaques
Tfie Cartfiaginian

down

general Hanno

the west coast of Africa

and the Periplus (account)


there

is

voyage

the 5th century BC,

in

of this

an intriguing reference

made

has survived,

in

it,

to wild, hairy people,

called 'gorillae' by local interpreters, living on an


island

in

a lake. Apart from this, the first convincing

Thomas Huxley," who reviewed

accounts and then current knowledge

from

whom most
Pigafetta's

Philip

Congo published

Description of
in

1598.' This

is

in

there

is

apes

is

was based on

look very

"in

Jones

in

1912.

delight to the nobles by imitating

much as

they are based on a faithful

if

description of a gorilla or chimpanzee.

Much more

and convincing des-

detailed

two books published

in

Purchas. He records the accounts given

it,

by the

brothers De Bry of two of these creatures, which

Andrew

the

sailor. In

this

a subse-

in

plate

is

in

passage that states (Huxley's translation):

Songan country, on the banks

volume

the

of

of

there are multitudes of apes, which afford great

the

modern sense. However,

the

in

quent chapter

following decades by the English clergyman

Kingdom

19th

assume from

1863 land

The Childhood of
Yarrow

least the

at

to

criptions are found

Animals by

E.

until

no reason

is

historical

drawn],

tfie

notes of Eduardo Lopez, a Portuguese

This 'group of young


primates' appeared

of the following

in

generalized fashion to

verbal account that the creatures referred to were

written record of man-like apes, at least according


to

[l^lacBca spp.),

century and there

First recorded contacts

in

World primates, particularly

Old

of

Battell, a soldier

Africa for

many

volume,"

who had

The longest

years.

to

him by

lived in equatorial
is in

the second

which Purchas recounts

in

the

in

Samuel

Battell's

description of forests along a river:

of the Zaire,

human

gestures".

Mary Evans Picture Library

Tl^e

woods are so covered

apes and parrots,


travails in

them

tfiat

is

the lesser

common

The

dangerous.

monsters

feare any

will

in

called Engeco. This

stature than a

in

man:

two

these

of

and

their language,

in

Pongo

proportion like a man,- but that he


giant

to

these woods, and

greatest

Pongo

called

is

man

alone, l-tere are also two kinds of

monsters, wfiich are


very

baboones, monkies,

witti

it

he

for

is

is

in all

more

like a

very

is

tall,

and

hath a man's face, hollow-eyed, with long haire

upon

his browes. His face

haire,

and

his

hands

but not very thicke; and

have no

but

hands clasped

He

of a dunnish colour.

is

in his legs: for

Hee goeth alwaies upon

calfe.

carrieth his

it

man

from a

differeth not

and eares are without

also. His bodie is full of haire.

in

the

his legs,

they

and

nape of his necke

when he goeth upon the ground. They sleepe in the


trees, and build shelters for the raine. They feed
upon

fruit that

they find in the woods, and upon

nuts, for they eate

no kind of

flesh.

They cannot

more than a
countrie, when they

speake, and have no understanding

The people of the

beast.

travaile in the
in

the night:

woods make
and

gone, the Pongoes


till it

in

the

wilt

where they sleepe

come and

sit

about the

fire

goeth out: for they have no understanding

lay the

There

wood

is

to

together.

an added marginal note from Purchas:

The Pongo a giant ape.

He

with him, that one of these

boy of his which

fires

morning when they are

lived a

told

me

in

conference

pongoes tooke a negro

moneth

with them. For they

"

Evolution, dispersal, and discovery of the great apes

hurt not those which they surprise at unawares,

He

except they look on them; which he avoyded.

was

their highth

twice as great.

a man's, but their bignesse

like

saw

What the other

the negro boy.

monster should be he hath forgotten to


these papers came to my hand since

my

which, otherwise, in

said

identical with

his death,

number

of

might

mentioned.

killers

was probably

but

of the ancients.

often conferences.

those

of

previously described by Battell, Tulpius, or Bontius,

pygmies

have learned. Perhaps he meaneth the Pigmy

Pongo

concluded that the animal was not one

and

relate:

time and

of the

Tyson reviewed existing literature

characteristics

"more resembled

"differ'd

which

in

so-called
large

animal

his

Man than Apes and Monkeys

do" and then a slightly shorter


it

the

He enumerated a

of

list

those

which

in

from a Man and resembled more the Ape

and Monkey kind." He concluded that though

The description
tarian habits

and building

of

nests

that of a gorilla, while the

phonetic version of Battell's

use

in

Gabon

down

of the 'pongo',

trees,

in

vege-

to its
is

clearly

name enche-eko'
was

engeco'l

at least until the early

la

in

still

9th century for

much resemble

"does so

more than any

parts,

animal'

do
-

in

look upon

'tis

of

A generation

later

can be found the

account

first

Europe. The Observationes Medicae

in

by Nicholas Tulpius, published

16A1. contains

in

may have been

young

chimpanzee, brought bacl< from the region

of

Angola

a description of what

to Frederick Henry, Prince of

and presented

Tulpius states that the animal concerned

is

a 'Brute-Animal sui generis'

is

this time, evidently, there

was already an
in

although confusion between

the various kinds persisted for well over a century


afterwards. This

is

Asia were garbled and

ridiculous.

ape'

The

Bontius

the

name

be seen there for

Buffon

pongo

finally

of

from

Battell,

the jocko,

Royal Society

in

London

in

saw
of

the

one

of

1699 and entitled, Orang-

outang, sive 'Homo Sylvestris', or the Anatomy of

compared

and a

'Man'.

chimpanzee, also

'Monkey' an

with that of a

The description

is

He thought

that the

small apes recorded by himself and Tulpius were

young pongoes.
Meanwhile,

in

1779 the Dutch anatomist Peter

Camper published

detailed

Orang, that

is to

on the

treatise

is

nor the Pigmy

of a

young

brought back from Angola.

specimen reached Europe;

first

in

the

Museum

seen by Camper

unsure

of

the

in

first
its

of

Tyson -

it

78^.

after

between

Battell,

tall

was

dis-

Orange and

Camper was

relationship

pongo

adult orangutan

skeleton

of the Prince of

animal, which stood over 1.2


called

is

neither the Pongo nor the Jocko,

of a peculiar species.

an animal

played

Borneo,

say, that of Asia, that of

of Tulpius,

first scientific

the

and a small one,

Africa;

the East Indies."

in

nor the Orang

account

Europe

many years. On

females and a young male.'^ He stated: "The true

have described an

17th century

Battell's

in

concluded that there were two spe-

which was indeed penned by

the

This,

orang', or manlike ape: a large one, the

...

of

from

adult ape recorded

Shortly after this, the

publication of the

'Ape'

last to

first

larj.

and Daubenton under

an Asian

the apes - a treatise by Tyson published by the

a Pygmie

be a lar gibbon Hylobates

the

posses-

into

gibbon from Asia

of a

basis of these specimens and existing accounts,

orangutan.

The very end

not only examined a

'jocko' (erroneously derived

was

engecol,

and the

in

consequently

may

1658,

in

specimen

sion of an adult

after the author's death,

really

who

orangutan, based on dissections of several young

rather hairy female human.

was added

but the description,

of

on the

often verged

apparent illustration

first

is evidently of a

This picture

a particular

have been the same

to

young chimpanzee, but also came

part explained by the fact

in

that, during the late 17th century, accounts of apes


in

French naturalist Button,


live

cies of

understanding that similar animals occurred


Africa,

It

detail by Buffon

western Sumatra."

in

seems

1739.

in

to

ape was not a chimpanzee from Angola, but an

both Asia and

and

animal later described and depicted by the great

described

Indies and southern Africa at the time, that this

orangutan from Angkola

appeared

(now known

possible, given Dutch contact with both the East

By

means

an undoubted chimpanzee

of

Orange.

the-Woods' and by the Africans as 'Quoias Monou'.


It

yet by no

accompanied by

description,

drawing by Scotin,

referred

Indians as 'Orange-autang, or Man-of-

to by the

of:

as the product of a mixt' generation

it

An anonymous
ape descriptions

Making sense
an ape

know

'species of Ape."

the chimpanzee."

of

it

of its

ape kind, or any other

of the

the world, that

many

a 'Man' in

evidently

this

large

and which he

and the

Ijuvenilel

orangutans he had described so meticulously


years earlier For

some years

it

was assumed

five

that

15

World Atlas

of Great Apes and their Conservation

it

was

chimpanzee,

a separate species from the

gibbon, and orangutan.

In

Blumenbach,

1810,

German, suggested the animal was


orangutan.' Richard

Owen

first

sl<eleton of an adult

an adult

monograph

also

recorded description of the

chimpanzee -

clearly the adult

version of the African animals considered by Buffon

and Tulpius

By

be young pongoes.

to

chimpanzee had

finally

become well characterized

as distinct entities, the former

living in

latter in Africa,

and both known

adult states.

was

other apes

in

It

Asia and the

their juvenile

in

and

also established that the only

pongo' -

described - Battell's true

lagged far behind.


In

1819 a traveler,

Thomas Bowdich, noted

that local people in the region of the

Gaboon (Gabon)

addition to the engeco, called the

described as

"five

feet high

and tour across the

shoulders."' However, there

evidence for the existence

when an American

ingena' and

was

of this

little

animal

missionary, Dr

further

until 1847,

Thomas Savage,

Mary Evans Picture Library

late 19th century print

unknown ape

the skull of an

Mr Wilson,

the Rev.

in

the

a missionary resident

From the

and the descriptions

skull

of the

animal provided by local people. Savage concluded


animal

that the

question

in

was

new species

of

ape." Savage and Wilson obtained a good account


of

the habits of this creature

anatomist,

description.

Wyman, an

It

publish

to

was Savage who

detailed

applied the

making no claim

species, although

actually the animal described

to

its

but, finally, the

account

of Battell's original

the African apes

was confirmed. As Huxley

the gorilla had of

all

being the

was known

apes;

gorilla.

it,

made known

to the

general

of the 19th century, therefore,

that there

were four

distinct 'kinds' of

eastern Asia, the gibbons and the orang-

in

utans;

of

be scientifically investigated."

last to

By the middle
it

put

the apes "the singular fortune

be

first to

world and the

being

the Periplus.

in

had taken over 200 years

remarkable accuracy

of

name

taken from the Periplus of Hanno, to this

gorilla',

It

the wild and enough

in

material to allow Jeffries

American

River reported the existence of a second great ape


in

of

on the Gaboon River

eastern Asia were various species of

gibbon. Understanding of the last of the apes to be


scientifically

house

physical

then, the orangutan and the

this time,

came across

demonstrated

finally

this in persuasive fashion.""^ His

contained the

in fact

in

western

Africa, the

Understanding

chimpanzees and the

of the natural history of the

gibbons and orangutans was quite well advanced,

thanks
lists

to the

number

observations of a

of

natura-

including MiJller, Duvaucel, Bennett, Wallace,

of the skeleton of a

and Brooke. There were known

human compared

species of gibbon, but the question of whether there

that of a gorilla.

with

were

several, two, or only one species of orangutan

was regarded by Huxley


Given

as unresolved.

at least

geographic

wide

the

area

chimpanzees occurred, he thought


there

be several

to

over

it

which

possible that

more than one species. The

might be

recognition of the bonobo as a separate species

the early 20th century confirmed this.

he assumed there

He also noted

be only one species

to

and Wilson, knowledge


the gorilla

in

accounts

that, despite the

the wild

of both the

was much

in

contrast,

In

of gorilla.
of

Savage

chimpanzee and

less

complete than

that of the Asiatic apes.

Apes as human

relatives

Although they were undoubtedly regarded as


fascinating

in

their

regarding apes

in

own

right,

western

intellectual circles at this

where they stood

in

the great question

scientific

time was,

relation to

and wider
of

course,

humans. Darwin had

published his revolutionary On the Origin of Species


in

1859, spurred on by Wallace,

who had indepen-

dently developed similar ideas largely as a result of

16

Evolution, dispersal, and discovery of the great apes

As evolutionary concepts

his observations in Asia.

took hold,

became ever

it

clearer that similarities

between species might be

were

more

the

affinities: that is

a sign

each other, the more

to

have shared a recent

evolutionary

of

similar different species

common

they were

likely

to

ancestor The notion

of the transmutability of species implicit in this,

although by no
to accept,

as

means new, was

was

difficult for

many

the idea of selection acting on

A century after

inheritable variation - the cornerstone of

random

evolutionary theory took

Darwinian theory.

There were enough problems


these concepts

when

but such difficulties paled

encountered
the

scheme

in

that the great

in

comparison

It

apes had

was

to

startlingly similar

and were thus closely related


-

anathema

who found

to

many

to

mean

one

of

those

most repugnant was Richard

this idea

most accurate and detailed descriptions

far the

to date of

the morphology of the chimpanzee (now considered


the closest relative to humansl. So determined

demonstrate the separateness

to

he erected an entire

Archencephala -

to

of

mammalian subclass

contain

them

human

brain, the
in all

of

others

was

who embraced

evolutionary theory

was

humans were

in

not whether

closely related -

taken as read - but which of the species

the closest

living

relative

to

humans. On

anatomical grounds, Huxley concluded that

it

was

either the gorilla or the chimpanzee, but believed

there
of the

was

was

insufficient evidence to

determine which

two was actually the closest. This question

not satisfactorily resolved until over a century

later,

likely to

highly

number

with the development of

new techniques

in

of a stick.

characters are

of

be related, and that species sharing

complex and specialized feature are

some

likely to

be more closely related than species not possessing


it.

Until recently, the features that could be analyzed

were

essentially anatomical or morphological ones,

although occasionally animal behavior has also

been used. There

are,

however,

difficulties with this

and, most importantly, because the

ramifications, the question

was

species sharing a large

evidently appear and disappear through evolution

number

San Diego Zoo

mashed end

hippocampus minor, supposedly

other apes.'

at

William H Calvin (www.williamcalvin comi

approach. These arise because characters can

in

way

perceive

soaks up juice with the

the

the great apes and


that

- the

alone, based very

To Darwin. Huxley, and the increasing

all its

was

humans that

largely on the presence of a small structure

absent

bonobo

each other was -and

Ironically,

humans

themselves. Here, a

anatomical

Owen, the anatomist who had produced by

he

that

and humans had shared a recent ancestor

that they

use among

revolutionized the

incontrovertible by

features to humans. The idea that this might

still is

of tool

great apes further

those

humans in
now

considering the place of

of things.

hold, the discovery

accepting

in

applied to other organisms,

ters can arise independently

Wings
in

for

birds,

powered

flight, for

mammals

(batsl,

in

same charac-

different lineages.

example, are present

and insects, and have

therefore arisen independently at least three times


in

the course of evolution. Moreover,

among

and insects there are species or groups


that can no longer
of

some

fly

and which may.

birds

of species

in

the case

insect groups, no longer possess recog-

nizable wings at

Advances

all.

in

molecular biology since the 1970s

have revolutionized approaches


systematics. The opportunity

now

to

taxonomy and

exists to

compare

molecular biology.

directly the genetic material of different individuals,

EVOLUTION OF THE GREAT APES

clearer insight

Reconstructing phytogenies: fossils and genes

between them. These techniques allow us

populations, and species, and often to gain a far

There are two major sources


be used

to establish the

of

evidence that can

relatedness of organisms:

the living organisms themselves and fossil remains.


In

both cases, two basic premises are used: that

mine which

into

living

the

degree

of

relatedness
to deter-

organisms are most closely

related to each other with increasing confidence.

However, describing the evolutionary route by which


these organisms arrived at their current state

still

17

World Atlas

of Great Apes and their Conservation

Box

WHAT MAKES A PRIMATE?

1.1

Together, the above features allow us to assign a

range

There
all

no unique feature that

is

primates, fossil and

be distinguished from

common
in

number

are a

there

other

in

living,

other

all

characteristic of

is

to

agreement

that

are

which

of

Rather,

is

not

found

be distinguished from other groups.'^


of

that the

thumbs

toes and

now widespread
all

share

more recent common ancestor with each other

than they do with the other primates. They therefore

form a monophyletic group, the simians. The

precise relationships between simians and other

primates, however, as well as the origin of the

these are:

feet that

is

monkeys and apes

primates as a whole, remain

hands and

the order Primates,

apes (including humans). There

mammals, each

The most important

to

by which they can

most primates; taken together, these allow

primates

animals

living

mammals.

features

of

of

Including tarslers, lemurs, lorises, monkeys, and

much

less settled.'^

can grasp, usually with big

opposed

that can be

Martin Jenkins

to the

other digits (although humans, for example,

have

the

lost

ability to

oppose

some

nail (although

one or two

form a

to

flat

bonobo's

foot,

with opposing big toe and

precision grasp.

species have modified

called 'toilet-claws', for

nails,

their big toesi;

claws that have been modified

William H. Calvin Iwww.wjlliamcalvin.coml

grooming on

and the aye-aye from

toes,

Madagascar and the New World marmosets


and tamarins have re-evolved claws from
nails on all digits except the big toej,

eyes that are at the front of the face and look


forward, with overlapping visual fields, allowing binocular vision

and accurate judging

of

distances;
relatively

compared

brains

large

normally found

in

other

those

to

mammals

com-

of

parable size;

small

litter

sizes - usually of only one young -

and young that mature slowly compared

most mammals
a

origin within

distinctive

to

of equivalent size;

the skull of the

auditory bulla (the bony case that protects the

underside

of

the inner and middle ears(.

requires evidence from the past, usually obtained

fossils are generally

from

of the

fossil

remains. The problem

is

sample

of life in past

Individual

organisms leave no

lasting physical trace

they die, largely because decomposing and

if

some

or

all of

efficient at their

an organism

chance that the remains

is

to

work.

recognizable

be recovered today

extremely small. Where they do survive, the

fossil

materials are usually very incomplete. Although

trace

fossils

may sometimes

give

exquisitely

detailed Indications of the soft tissues of organisms.

18

formed only from those parts

organism that were hard

in life

- for example,

shells of mollusks, bony skeletons, and,


of

many

cases

vertebrates,

teeth.

Only

of recent preservation in the

remains

is

in

form

in

the case

exceptional
of subfossil

there any likelihood of genetic material

being recovered.

preserved, the

will survive in

form through geological time


is

biased

times. The vast majority of

scavenging organisms are so

Even

that the fossil

an extremely incomplete and

record

when

is

Overall,

It

Is

estimated that known fossils

represent perhaps

have ever existed." For the primates, the


record

may

percent of the species that

has been estimated

to

be rather

fossil

more

complete, at up to 7 percent representation."

The geographical clustering


yield

of

fossil

finds

can

even more complete series than this overall

Evolution, dispersal, and discovery of the great apes

would suggest.'^

figure

Still,

any one lineage,

in

we

nnany more species have existed than

know

among

about, even

the primates, which are

and studied.

particularly intensively searched for

One

corollary of this

that

is

any known fossil organism


of

any

it

very unlikely that

is

the direct ancestor

is

one. This simple observation

living

often overlooked

in

human

to recreate the

Most recent attempts


dar

more so than

ancestral

in

which bring together molecular studies and the

They are based on the assumption

fossil record.

random mutations accumulate

DNA

that

various kinds of

in

constant rates, with no reverse mutations,

at

and that code differences between lineages can be


calibrated with the fossil record and
lent to

made

equiva-

time since lineages became separate.

DNA le.g.

mitochondrial

DNA or mtDNAl

appears

to

the globin genes), so their molecular clocks

can be said

run at different rates. These different

to

rates can be exploited to inter evolutionary rela-

tionships at different taxonomic levels,


less far

used

back

human

in

more

mtDNA

time. For example,

can be

populations, globin genes for relationships

between modern mammals, and cytochrome


relationships
lineages.

The reason why the cytochrome

molecule

c for

deep time' between eukaryote

in

runs so slowly
this

or

between modern

establish relationships

to

i.e.

is

that the structure

is

vital

to

clock'

and function

of

metabolism and cannot

be changed through significant mutation without


lethal

These molecules therefore remain

effect.

essentially the

same across

very

many

measure

years requires the

of

one or more

this

to

ancestors

to

form two

superfamilies

the

species,

of

groups were prosimians

Box

1.11,

a range

northern hemisphere. These

of fossil sites in the

(i.e.

and some 200 species

simians, see

not

over 70 genera

in

have 50 far been described.^' The vast majority


of

them had disappeared by the end


from the

(at least

fossil record),

Eocene

of the

apparently falling

known from

an actual

fossil record to offer

known

the fossil record, only nine are

from post-Eocene deposits." The body sizes


these early primates are estimated

from around 50 g
primates: the

up

(the

weight

of the

of

have ranged

smallest

mouse lemurs, Microcebus

7-8 kg (the weight

to

of the

to

living

spp.) to

larger guenons,

Cercopithecus spp., or the smaller mangabeys,

Cercocebus

spp.).

By the early Eocene, the primates were


already evidently well established as a group,
indicating that their origin lay earlier, at a time for

which no relevant

fossils

how much

earlier

Exactly

Until relatively recently

it

have yet been found.


is

a subject of debate.

was widely argued

the temporal origin of primates, along with

other major
to

mammalian

that

most

groups, lay relatively close

Cretaceous/Tertiary boundary. At around

the

65 mya. this

is

the point at which the dinosaurs,

many other groups


was assumed that the

plesiosaurs, pterosaurs, and

disappeared.

finally

It

mam-

primates and others diverged from early


malian stock

at this

time and then underwent a

relatively rapid radiation, evolving into a


of

forms over what

is,

wide range

geologically speaking, a fairly

short time.

More recent analysis, however, using molecular


" and applying statistical analyses to the

species, but

nevertheless accumulate minor differences from

harmless mutations. Relating

groups

Omomyoidea and Adapoidea, recorded from

Some

accumulate mutations more quickly than others


(e.g.

common

from

radiated
distinct

prey to deteriorating global climates. Of the species

line.

to recreate the calen-

use molecular-clock methods,

evolution

of

very

is

debates about genealogy and

phylogenetics, perhaps nowhere

attempts

currently

clocks^'
fossil

record."'

suggests that divergence

of

the

major mammalian groups, including primates, can

reliable estimates of divergence time in

the lineages under study. Both the assumption of a


regular rate of change

in

DNA through

time and the

Table

1.1

Epochs

of the

Cenozoic era, the 'age of mammals'"

setting of calibration points are problematic, but


this

that are gaining

case

Start (mya)

approach has nevertheless produced results

of the

wide general acceptance, as

in

the

apes discussed below.

Primate origins

The earliest fossils that are unequivocally identified


as primates date from the early part of the Eocene

epoch (Table

1.1],

some 5A-55

mya.^'

"'

They rep-

End (mya)

Paleocene Epoch

65

54

Eocene Epoch

54

38

Oligocene Epoch

38

26

Miocene Epoch

26

Pliocene Epoch

1.6

Pleistocene Epoch

1.6

0.01

Holocene Epoch

0.01

now

resent a diverse collection of lineages that have

19

World Atlas

of Great Apes and their Conservation

be traced back

much earlier, to the mid-Cretaceous


some 90 mya, with the last common ancestor of

some

era

primate was an already arboreal animal that

all

the living

primates believed

have lived

to

somewhat over 80 mya.


Eocene primates

is

highly problematic,

and

ideas about their hypothetical ancestors are even

more

many years

speculative. For

it

was argued

that

was

the

the major impetus for primate evolution

adoption
ing

of

an arboreal

hands and

lifestyle,"

for

feet

"

holding

size

for

ground). There

is,

in

however, one major problem with

perse

the chimpanzees of

primate adaptations: the fact that

characterized and
to

known

be quite distinct from

are at least as arboreal as

mates and

yet do not

Southeast Asia.

was

likely to

possess these attributes, or

at

them together"

More recently

the orangutans of

explains

many other
many pri-

mammals

least not all of

(of

windy canopy above the

the notion that an arboreal lifestyle

were well

it

flowering

the

of

species

range

has been argued that

diet

Michael Huffman

order

in

to

(angiospermsl.

the Eocene coincides with a

in

range

In

primate

of

marked

flowering plants and particularly the

in

complex

of

tropical forests, with a

They argue that primates

of fruiting trees.

evolved specifically to take advantage of this

new

and flowers growing

of nutritious fruits

in

the fine, terminal branches of trees and bushes.

Studies of

modern lemurs such as

M/crocefaivs and

the dwarf lemurs, Cheirogaleus, believed to be


quite similar

in

many ways

suggest that the spur


primate features

to

these early primates,


developing fullblown

to

may have been

the need to adapt

combination of these two food sources. The

to a

have been a principal impetus, with

plants

apparent radiation

the

particular,

development

By the early 19th century,

Africa

rise

onto branches,

processing complex spatial infor-

hunting by sight

the fine branches of the forest

bushy undergrowth. Others note

in

radiation

mation, and a decreased dependence on smell


lesser importance

canopy or

to

in

that the rise of primate species coincides with the

involving grasp-

binocular vision forjudging distances, an increased


brain

became adapted
capture insects

Reconstructing the ecology and behavior of


the

arguing that the ancestral

primatologists

fruits

and flowers on the

fine

terminal branches

would themselves have attracted


insects and

range

of

other invertebrates, and the early

primates would have fed on both the plant matter

more

and, perhaps

opportunistically, on the con-

gregating invertebrates." As will be seen, diet and,

changes

particularly,

in

diet are widely held

have played a crucial role

primates from the origins

appearance

the

in

of the

to

evolution

group up

of

to the

modern humans.

of

The

origin of the simians

The

earliest evolutionary history of the primates

still

a mystery, as

emergence

of

is

the

is

the exact path that led to the

simian

line.

Possible fossil

now known from as far back as the


Eocene, some 50 mya, from a range of sites in

simians are
early

North Africa, although most Eocene simians are

from the

Eocene, and the earliest known from

late

outside Africa (from the Arabian Peninsula] date

from the early Oligocene. The relationship


early simians to the

unclear -

it

far

is

from certain that either

two groups actually gave


IS

it

the

clear
initial

made

in

deposits

these

of

is

these

rise to the simians.

where or when the simian

Nor

line arose.

As

discoveries of early simian fossils were

North Africa (specifically the Fayum


in

Egypt],

simians arose

in

possible

they

that

subsequently

20

of

Omomyoids and Adapoids

it

was widely assumed

that

it

is,

however, perfectly

arose

in

Asia

Africa;

to Africa.

and spread

Evolution, dispersal, and discovery of the great apes

Old World Simians and the Miocene radiation

known from

Fossils are

the late Eocene and early

Ollgocene, 30-^0 mya, that are indisputably early


'catarrhine' primates, of the lineage that includes

both the apes Ithe hominoidsl and the Old World

monkeys

(the cercopithecoidsl.

been found

Taqah sediments

Most

Fayum deposits

the

in

Oman,

in

in

but there

these have

of

Egypt and the


is

also a single

tooth from Angola." These fossils, of which the best

known

is

Aegyptopithecus, provide a link between

the early Eocene simians and the living

and apes. The nature

of the fossils,

monkeys

and molecular-

dawn

clock analysis, indicate that these so-called

apes pre-date the period

at

which the apes diverged

from the Old World monkeys.

The Miocene - the age

the apes

of

For the Old World primates, as indeed for


other groups

many

animals, the Oligocene represents

of

an important gap

in

the fossil record. During the

Miocene, around 22 mya, however, fossils

early

begin to reappear

in

much

greater numbers. By this

were evidently

time, the apes

firmly established as

a separate evolutionary lineage, indicating a split

with the Old World

monkeys some time between

22 and 30 mya. The Miocene, which ended around

seen as the era

5 mya, can be

which

of the

apes, during

became remarkably widespread

this lineage

KOCP

and diverse

the Old World.

in

perhaps 40 genera have so

numerous
Research

locations
in

in

Up

100 species

been

far

Africa,

to

in

identified at

Europe, and Asia.

and the rate

this field continues,

new fossils indicates that


species named so far constitute only a

many

the

discovery of

of

fraction of

lack characteristic of

appears
of

to

However,

life.

A new

branches, and
along the tops

was
in

reported

Spain.^^

in

Many

from only very

200A from mid-Miocene deposits


of the finds,

partial

however, are known

remains -

chiefly teeth

associated fragments of jawbone - making


difficult to

it

and
very

determine phylogenetic relationships or

reconstruct the adaptations and ecologies of the

Known ape

fossils

from the

first

part of the

Miocene, until 15-17 mya. are confined to Africa.

From what can be determined,


early

the

Miocene apes,

of

genus Proconsul, were

ranging

in

it

seems

that these

which the best known are


a

variable

in

group,

probable body size from around 3 kg to

had not developed the highly

It

seems

branches on

that

with a range of other


of a

new

apes, that

suited to traveling

Orangutans are believed


to

have diverged from

the

common

ancestral

line of the great

some

apes

11 million

years ago.

all fours.'

15 and 17

Such

mammals,

taking advantage

land bridge between the two continents.

land

had

bridges

migration, however, so
in

it

before

existed

this

something

that

is likely

the biology of the apes (or the

ecology of Eurasia!

to

allow them

to exploit

and

occupy Eurasian ecosystems: perhaps a dietary


factor which enabled

them

herbage and/or harder

to subsist

fruits.

on coarser

Fossil

evidence

indicates rapid dispersal and diversification, with

apes being widespread and diverse


record from around

frugivorous, although diet undoubtedly varied from

mya.^

In

modern

some time between

fossil

tail

of

was thus more


of

well over 80 kg. They appear to have been largely

species to species. Proconsui at least lacked a

and

mya, the apes invaded Eurasia from Africa along

had changed

species concerned.

an arboreal way

to

enable them to travel by swinging along beneath

genus and species, Pierolapithecus catalaunicus,

the likely true diversity of Miocene apes.

it

powerful forelimbs

flexible,

modern apes

all

have been adapted

in

14

in

mya

the Eurasian
to

around 8

contrast, the fossil record for large apes

Africa for this period

is

very sparse. African

21

"

World Atlas

of Great Apes and their Conservation

now

is

widely accepted that the gibbons were the

to

first

split

from the branch that has

off

led to

the other living apes. The orangutans are the next

have diverged. Gorillas are believed

to

to

have

split

next from the line leading to chimpanzees, bonobos,

and humans, with chimpanzees and bonobos being

most recent

the

from each

to diverge

other.

Although the sequence of events


disputed, the timing of each split

open

to debate. This is

reliable calibration

assumptions
in

the

that

because

from the

DNA sequences
of a

depends both on

it

at 23.3

and on

fossil record,

One study

that are analyzed.

range

of

sequences, and uses

date of divergence between Old World

apes set

no longer

much more

uniform rate of mutation

of a relatively

makes use

is

is

monkeys and

mya, gives divergence times as

follows (with 95 percent confidence limits!: gibbons


14.9 2 mya; orangutans 11.3 1.3

6.A

5.4 1.1

number

mya; gorillas

mya; and chimpanzees and humans

1.5

mya." These generally agree well with

although the divergence

of earlier studies,

times for the gibbons and the gorillas are somewhat

more recent than those

given

Pilbeam, which are 17-16

mya and around

tor

by,

assumed divergence time

World monkeys and the apes back


Oligocene would

times

of

mya

Changing the calibration point by

respectively."
shifting the

example,

the Old

of

into the

late

the calculated divergence

shift

the various apes back, but the order of

divergence and degree

of

relatedness between the

species would remain the same."


Martha

The

gorillas, while the

first to

be recorded,

were the

last of the

remains from around

fossil

Robbins

mya onwards, which


more

consist almost entirely of apes believed to be

or less closely associated with the hominid

much

great apes to be

are

scientifically described.

undoubtedly
effort

better represented

spent

From
living

in
in

is

it

hard to

and

living

relationships

it

is

of

former ape

of the
is

descent

so scanty.

ape fos-

speculation as to the exact


living

that,

most frequently

in

the case of

Asia have been associated with this lineage, most


clear that the

of the l*/liocene

between the

do just

the orangutan. Several genera of fossil apes from

species (including humansl. These

much

ones

to

also very hard to demonstrate close anatomical

factors have led to

to living

is

make sense

between any

apes

of

searching for them.

is

difficulty of linking fossil

has not stopped paleoanthropologists from trying

amount

species since the fossil record

relationships
sils

line,

Possible orangutan relatives

The

collections; this

apes represent a small vestige

of living
It

in

part because of the large

the fossil record,

diversity, but

species,

and

importantly Gigantopithecus from


deposits

in

and Pleistocene deposits


Viet

late

Miocene

the Siwalik region of India and Pakistan,


in

southern China and

Nam; Lufengpithecus from

and possibly early Pliocene

in

the late Miocene

southern China;

Sivapithecus from the late Miocene 112.5-8.5 mya)


in

the Siwalik region; and Khoratplthecus from the

middle and

The

late

Miocene

latter three

in

Thailand.

have been proposed as close

regarding the nature of their immediate progenitors

relatives to ancestral orangutans, although strong

and where they may be found.

arguments have also been made against

Some

of

this

speculation - at least that

concerning the relatedness

now been

22

of

the

living

apes - has

largely resolved by molecular analysis.

It

this in

the case of Sivapithecus and Lufengpithecus"'

Currently there

Sivapithecus

is

seems

to

be agreement that, while

not particularly closely related to

Evolution, dispersal, and discovery of the great apes

the ancestral orangutan,

orangutans than
however,

is,

to

much

less

closer to

least

at

is

it

any other

living

primates. There

consensus on whether

this

The new genus Khoratpithecus (which includes


two species,

of

which one,

originally placed in

seems

at

some

share

chiangmuanensis, was

Lufengpithecus vjhen described!

present to be the best candidate for a near

orangutans,

fossil

with a tropical flora. This

also

it

have

associated

is

have

lived

in

areas with temperate or

seasonal and relatively open rather than forested

dated

Khoratpithecus species,

Miocene

(just

over

with a flora that


indicating a

11

earlier

the

Interestingly,

mya),

association

shows strong African

temporary

floral

middle

the
in

two

the

of

to

found

is

affinities,

and faunal dispersal

corridor between Southeast Asia and Africa at this


time. This

gence

fits

(11.3

well with the estimated time of diver-

myal between the orangutan lineage

and that leading

to the gorillas,

chimpanzees, and

humans, as derived from molecular-clock


the gibbons represent the survivors of the
of

analysis.

might speculatively be argued from this that

It

apes

to

genus was con-

surviving into the Pleistocene, this

temporary with

in

deposits

in

to this

humans; indeed,

early

least

at

species have been found

i^omo erectus and orangutans

of

Vietnamese cave dated

latter

tribe (the Sivapithecini!." In

around

to

475 000 years ago. The timing and cause

of

its

extinction remain, tantalizingly, a mystery.

contrast to both

in

is

comes
to

Sivapithecus and Lufengpithecus, both of which

habitats.

same taxonomic

the

mixed with those

occurred during the Pleistocene, and

to

in

the

line,

and has been tentatively grouped with the

remains ascribed

from an area where orangutans are known

human

to the ancestral

be most similar to Sivapithecus

to

orangutan." Not only does

and Pleistocene

appear

or dead. Although at one time thought

living

be perhaps close

highly distinctive features with living

relative to the ancestral


it

K.

known,
to

genus appears

the case for Lufengpithecus.

is

(mandibles!, indicate an ape larger than any other

first

wave

invade Eurasia during the early Miocene,

while the orangutans represent the survivors of a

Chimpanzees and humans

much

While

the

in

evolution

the

of

apes

great

remains uncertain or strenuously debated,

it

is

now

almost universally accepted among scientists that

chimpanzees are our nearest


split

leading

and

A.3

relatives,

between the lineage leading


to

6.6

modern humans took


mya, and that

last

common

ancestor

earliest distinct

all

of

that

between

human and

in Africa.

What the

two groups and the

of the

hominid (and indeed the ancestral

chimpanzee! looked

why

place

phase

this

chimpanzee evolution took place

that the

them and

to

how

like,

they behaved, and

the two lineages went their separate ways are

the subject of

much

speculation. Characteristics

that are widely considered to be very important

distinguishing the

human

in

lineage from that of other

apes, and particularly the chimpanzee lineage, are;

subsequent, mid-Miocene, invasion from Africa.


Alternatively,

African

apes,

including

secondary invasion
apes,

some time

hominids, arose from a

of that continent

living

of this

larger

two Eurasian genera, Dryopithecus, known from

be close

to

the ancestor of African apes and humans.' There

is

currently no consensus on this, and

it

is

unclear

how

in

an

humans than

in

in

is

far

any other primates;

and
tooth structure and wear, which

Spain to eastern Europe, or Ouranopithecus (or


likely to

the only

brain size relative to body size, which

hypothesis regard one

Graecopithecus] from Greece, as

modern humans being

hominids that habitually engage

erect, bipedal striding gait;

by Eurasian

during the middle or late Miocene.

Modern proponents
of

bipedalism.

has been argued that the surviving

it

correlated with

Great importance

is

is

strongly

diet.

attached to teeth because these

such a consensus might be reached, although an

are the most abundant, and sometimes the only

the fossil record, particularly of mid-

available, fossil remains. Interpreting tooth struc-

improvement

in

and late-Miocene African apes, would help.


Gigantopithecus, the other probable orang-

utan relative,

is

unique

in

nonhominid ape genus so


vived

later than

being the only extinct


far

known

the early Pliocene.

to

have sur-

Pleistocene

ture from often

evolutionary
Similarly,

bipedalism
the

human

worn and broken

arguments

although
is

it

is

is

fossils to construct

highly

contentious.

widely accepted

that

fundamental feature distinguishing

lineage from that of other living apes,

no consensus as

when

arose. There

remains from southern China and Viet Nam, as-

there

cribed to the species Gigantopithecus blackii, and

are currently too few relevant fossil remains to

consisting of extremely large teeth and lower jaws

construct convincing arguments. Brain size

is

to

it

is

of

23

World Atlas

of Great Apes and their Conservation

limited use

constructing early phylogenies as the

in

major increase

in relative

brain size

began only around 2 mya, long

line

the

in

human

after the split

from the chimpanzee lineage.

may have

played an

Currently there are three major candidates for

human

lineage.^"

important part

perhaps

evolution, related

the earliest fossil of a distinctly

These

These enzymes are associated with a meat-

lineage.

eating diet and indicate that increasing carnivory

authors agree that

final point is that not all

genetic divergence has yet occurred

sufficient

between chimpanzees and humans

are:

being placed

Ardipithecus kadabba from Ethiopian deposits

dated at 5.6-5.8 mya;"

"

separate genera."

in

were accepted, then

warrant their

to

this

If

taxonomic

for

argument
purposes

chimpanzees, bonobos, and humans would

Orrorin tugenensisfrom deposits estimated to

date from slightly under 6

mya

in

the Tugen

Kenya;" and

Hills in

human

in

to increasing brain size.

assigned

to

the genus

be

all

Homo, further emphasizing

the sibling nature of our evolutionary relationship


with these great apes.

Sahelanthropus tchadensis from the Durab


Desert

northern Chad, dated to between 6

in

RESEARCH ON WILD GREAT APES


The

and 7 mya.'

Each

these (very incomplete) remains appears to

of

have a mosaic
that
to

of so-called primitive characteristics,

those shared with earlier fossil apes and

is

some

extent with

characteristics, that

apes, and derived

living

those more closely aligned

is

with the hominid lineage and not shared with other

apes. Each has

living

own

fervent supporters as

known representative

the earliest
lineage,

its

and each has

detractors.

the

If

its

own

is

accepted

15. i

1.1

chimpanzees and

myal," and the dates

and Sahelanthropus are

for Orrorin

human

most recent molecular-clock

analysis for the split between

humans

of the

equally fervent

reliable,

then

who spent weeks

at

London Zoo watching

monkeys before he wrote Expression of the

the

Emotions

in

Man and Animals

primatology began

chacma baboons [Papio

century, with the study of


ursinus]
this

South Africa by Eugene Marais, though

in

was

not

published

long

until

afterwards.'"

Primatology continued with attempts by Henry


Nissen^' and Harold Bingham'' to observe chimp-

anzees and gorillas

in

the wild - described by Alison

Jolly as "difficult quarry in impossible terrain, for

who had no

people

what primate research

idea

would mean."'" Clarence Ray Carpenter accom-

monkeys

plished field studies of howler


palliata]

mulatta]

in

ancestors maintain that the molecular clock

flower

the 1950s and 1960s,

is

in

the 19^0s.'^ Field primatology began to

when an

the study of the great apes

earlier than

Japanese researchers were among the

indicates.

it

dence

for the

fossil

record provides scant evi-

changes

that took place leading to the

the chimpanzee and

divergence

of

some

might be shed on

light

the genetic material or

genome

of

this

human

lines,

by comparing

DNA code that comprises the

each species. This

is

for the current undertaking to

field,
in

chimpanzee genome. The genomes

of

Kortlandfs

field

trips

One

Early

research focused on eastern chimp-

humans

Schaller pioneered studies of mountain gorillas

some insight into what, at least


makes us distinctively human."

interesting preliminary finding

for breaking
of proteins!

human

down amino

is

that

enzymes

acids (the building blocks

have been positively selected for

lineage,

Belgian

the Congo)

anzees, mountain gorillas, and orangutans. George

in

at the genetic level,

of

the

fully

hoped that examining the remaining

percent will give

the

from 1960 onwards."

followed up by Dian Fossey

1.2

then

the

to

Congo (now Democratic Republic

same, and

is

first in

with an exploratory trip by Imanishi and Mitani

and chimpanzees are around 98.8 percent the


it

interest in

the wild emerged.

in

1958. Dutch researchers followed soon after

with

one major impetus

sequence

[Alouatta

and lar gibbons [Hylobates lar] in the


1930s,'' " and of rhesus macaques \Macaca

wrongly calibrated and that divergence times were

While the

Field

(18721.

the early years of the 20th

in

may pre-date the time when the


chimpanzees and humans diverged. However,
those who advocate either of these as early human
these two at least

24

'primatologisf might have been Charles

first

Darwin,

in

the

compared with the chimpanzee

the 1950s and

1960s;""

in
in

1967, these

were

Rwanda, leading

to

the establishment of the Karisoke Research Center

where she worked

until

her death

Goodall's study on eastern

in

1985." Jane

chimpanzees began

1960, and led to the establishment of the

Stream Research Center


Tanzania.'" Both

in

in

Gombe

the United Republic of

women began

their studies

under

Evolution, dispersal, and discovery of the great apes

renowned paleo-

the direction of Louis Leakey, a


anthropologist. and their

work helped

significant international

first

apes,

through

particularly

to create the

awareness

of

great

pages

of

their

the

principal sponsor's magazine. National Geographic.

The

killing

animals,

of

one

Digit,

the

of

Karisoke study

1977 shattered the Eden-like quality

in

those early studies. The

Corporation TV series Life on Earth,


in

of

Broadcasting

British

screened

first

1979 and featuring David Attenborough face

to

face with mountain gorillas, and the sensational

murder
life.

Dian Fossey (followed by the film

of

world

mountain

to

at

Stream continues

to date.

The
began
in

first

work

Borneo,

central

in

again with the involvement

Camp

field site is called

at the

orangutan continued

same time

Leakeyl; this project has

Sumatran

earnest,^ "'^ followed by the

in

was

in

the

survey of

distribution

the northern range of the bonobo


1973,''

of the

the early 1970s," at about

A systematic

'

in

Louis Leakey (her

of

and the central chimpanzee

gorilla

1980s.^'

1971,'

research on the western

that

chimpanzee began
western

in

"

Tanjung Ruling

Indonesia

continued to date. Detailed studies

in

started

Sabah, Malaysia and Renun, Sumatra."

Reserve

out

Gombe

orangutans

when John MacKinnon

Birute Galdikas began to

the

her

perils facing

Karisoke and

both

significant field study of

the 1960s,

in

and the

gorillas

them. Research

out

of

Gorillas in the Mist], also helped to alert the

first

carried

and the threats from logging and

Martha M. Robbins

primate biologists, philosophers, and others


recognize great apes and
single

'community

humans

of equals', with

to

as belonging to a

common

Uganda.

in

has continued

to

and concern

Public interest

animals

deepen, and research on these


increasing at universities and

still

is

apes

for the great

scientific institutions

Some

throughout the world.

research centers have also played an important direct


role in conservation

Ketambe

in

of the great

apes, including

Camp Leakey in Borneo,


Rwanda, and Gombe in Tanzania. By
Sumatra,

attention

captive

in

the wild

was

studies,

in

carried out.
parallel with

and the disciplines asked and

answered questions

of

ape behavior, learning

which had been used


laboratories,

ability,

for

were the focus

research, though they

Karisoke

ground.

in

and deterred poachers and loggers on the

In

some

cases, the areas around the field

stations have been declared national parks.

each other regarding great


ecology, evolution,

and communication.''^ Chimpanzees,

cognition,

decades
of

most

became

in

biomedical

early behavioral

less prominent as

RESEARCH ON CAPTIVE GREAT APES


A

population of captive animals allows noninvasive

research
impossible

to

be conducted

in

interactions,

that

is

or

difficult

the wild, including studies of social

animal

health,

and

reproductive

humans share many

studies of other great ape species multiplied. The

biology Great apes and

discovery

features at the levels of whole-body physiology and

be taught

in

the early 1970s that great apes could

human

sociobiology

in

sign languages, and the birth of

the late 1970s, both contributed to

intense interest

in

The discovery

of

great ape behavior and evolution.'"'


culture,"'

recognizable politics
revolutionized the

the wild,

freedom from torture and arbitrary imprisonment."

research on bonobos

research developed

in

Bwindi National Park,

their simple presence, they have attracted political

Field

observed

rights to

bushmeat hunting were well understood," but


was not until the 1990s that further significant

it

Mountain gorillas being

tool

use,'

and even

among chimpanzees^*

further

way we humans thought about

great apes and defined ourselves

in

relation to

them.

By the 1980s, a movement had begun among

metabolism, organ function,


even gene organization;

this

cell

means

are excellent models for studies of

basic

structure,

and

that great

apes

human

health

and disease. These range from field-based studies


of

disease

in

the wild, to laboratory studies of

captive animals, and even the involvement of apes


in

space research.

fruitful

avenue

for research with

apes

is

25

World Atlas

of Great Apes and their Conservation

Box

1.2

CRYPTIC APES

gination guided by expectation based on legend can


interact with observation to confuse the issue

Cryptic species are those that are reported anec-

from physical signs Isuch as

dotally, or

footprints),

absence

presumed

the

mountains and

habitat of high

yeti

the

in

On the other hand,

hard evidence.

of

or from photographic evidence, but that have not

dense Rhododendron forests

been described unambiguously from a

or dead

conducive to observation, camera trapping, or the

The great ape

hunting and capturing of shy and elusive animals.

specimen by

a professional biologist.

community has

attracted

live

own share

its

of cryptic

species, of which three desen/e a mention

volume because some primate


that there

is

day give rise

new validated

These are the


pendek'

of

may one

species or subspecies.

the

of

'yeti'

this

biologists believe

a significant chance that they


to

in

Himalayas,^^ 'orang

Sumatra.'" and 'Bili-Bondo ape' of the

northern Democratic Republic

Congo IDRCI.

of the

remote areas

in

not

is

Following an expedition to Nepal, the mountaineer

Messner concluded

Reinhold

that

Himalayan black bear [Ursus


bear rears onto

the

front

of

Messner also

indicates that

see a

yeti,

belonging

was

it

is

The

when

foot in the footprint

which gives the tracks the

foot,

appearance

to

yeti

hind legs at times, and

its

moving often puts the back


of

the

thibetanus].^'

bipedal animal.

to

when

villagers took

him

bear that was encountered.

Yeti

There have been many reports by western, Chinese,

and other

Tibetan, Nepali,

bipedal ape-like beings

local obser^'ers of large

the Himalayan Mountains

in

Sikkim, Bhutan, and Nepal, and nearby ranges

of

such as the Pamirs

was that

published report
representative

Nepal,

in

The

Tajikistan.

of

of B.H.

in

1832.

earliest

Hodson, the

Many

British

reports since

have been by mountaineers participating

1921

expeditions to the

Mount Everest

in

area, variously

including observations of the animals themselves

Don WhiUans

1949.
1

Cameron

by Alan

(e.g.

Norgay

in

1953,

of

Dyrenfurth

sometimes

yeti's

in

Hillary

other

name

889,

and Tenzing

in 19781,

19581,

in

inhabit this part of the

in

le.g.

1954, and

The animals are

in

some

Hodson

in

1948, and the

19571, giving rise to the

A team from Fauna and


led

by Debbie

Martyr,

Flora International,

has analyzed completed

questionnaires from about

200 Kerinci

these support the original Dutch claim


that

IS

not a gibbon or orangutan as

All of this

19381.

anecdotal and fragmentary evidence

shows how myth and ima-

people;

an ape

of

we know them.

Apart from numerous casts of a unique footprint,

each team

and

IS

member has seen an

totally

convinced of

its

Holden.
in

so

orang pendek,

existence, despite

managed

in

was

claimed by the Dutch around 1920,

the current lack of proof from the

by d'Auvergne

suggestive, but also

evidence to suggest that the southern

helped by them

(e.g.

it

Sunda shelf? The modern

populations of Kerinci-Seblat are different, as

sometimes

abominable snowman', but

of the

is

orangutan has spread southwards around the

there are also reports of injured climbers being

IS

camera

successfully otherwise

by

traps,

Jeremy

Achmad Yanuar has surveyed orangutans

Borneo, and was very skeptical about the orang

the investigation of the aging processes. Because

that

apes are so much

behavioral change associated with brain aging. The

many

of

the

from many

of

like

humans, they go through

same aging processes and suffer


same age-related disorders as do

the

humans. Thus studies


explored

26

in

food scraps and dung

1889, Jan Frostis

Slick expedition

the canopy? Or

in

ancestral orangutan that used to

IS

Wadded

said to injure people (e.g. by

1832, Waddell

Thomas

in

agile gibbon

of the

faunal barrier of Lake Toba to the west," but there

by the London Daily Mail expedition

Norman

Is

in

and Lord Hunt

supported by findings

siamang and
descendant

person).

(little

has escaped competition with

in

Edmund

1936,

in

gibbon that

it

west-central Sumatra,

in

as 'orang pendek'

locally

ape

the Kerinci-Seblat National Park

and surrounding areas

known

ground-dwelling

bipedal

to exist in

Norgay

9981 or their footprints le.g. by L.A.

H.W. Tilman

undescribed

appears

and Craig Calonica

1923, Tenzing

in

1970,

in

Orang pendek

An

of

menopause and

aging

in

great apes have

the behavioral changes

accompany

Great

it,

changes

Ape Aging Project

in

sexual

involves

activity,

and

noninvasive

monitoring of health, cognition, and behavior of


the oldest great apes

in

research

zoological gardens. Further,

it

facilities

and

promotes the study

Evolution, dispersal, and discovery of the great apes

pendek,

human
that

it

and

saw

he

until

to validate

one.

rennains a ctiallenge

It

existence of

ttie

lineage; meanwhile, there

poaching and

must be pushing

this

the brink of extinction.

The context

speculation about the orang

of

pendek was dramatically changed


publication

the

by

fires

felling that are devastating

Kerinci-Seblat National Park


to

the real fear

is

soon become extinct because the

will

illegal

ape

species sharing

ttiis

fioresiensis. a

tall

lineage) that existed

anecdotal evidence of
times.''

''^'

October 2004

hominin

of

Homo

Ifossil of

human

on the Indonesian island

as recently as

Flores

in

descriptions

of

18 000 years
survival

its

ago,

into

of

This cast of the skull of the 'unknown ape' of Bili-

Bondo

This extraordinary discovery encourages

being used repeatedly and were often


locations,

reported, small, current, bipedal, ground-dwelling

Sumatra. One obvious

'ape' in

Homo

possibility is that both

erectus whose fossils dating

to 1.6

mya have been found on

which

lies

the island of Java,

between Sumatra and Flores.

which

North
1

000

of northern

Congo

of the

The following

and
to

Ammann

resemble

anzees
in

km between
This gap

gorilla].

in

had

of Bili

in

in

of

about

distributions of the

who measured

to slip

characteristic of
in

DRC

male

inhibited

scientists

Richard

in

captivity,

2003
skulls

be those of exceptionally

were also longer than the

of footprints

of a

dead individual exceeded that

of

the

recorded chimpanzee. During the 2003

gorilla

survey, Williams reported four of the apes being

attracted to an imitation of the cry of a

sighted a gorilla

local great apes.

These

duiker; they

approached

deadly intent before fleeing silently on encountering

humans. This behavior

away from people and

only opportunistic,

the intimidation display

chimpanzee than

gorillas.

The 1999-2002 war

team

of

including George Schaller and


visited the area

in

2001 and

apes that have aged and died

with a view to discovering

more about

in

the

wounded

and apparently with

fast

is

suggestive of an active,

hunting animal,
a

more

Mitochondrial

gorilla.

analysis of hair and fecal samples had

like

if

DNA

meanwhile

established a chimpanzee identity for the animals,


at least

on the maternal

University of

Researchers from the

side.

Amsterdam launched

these unusual chimpanzees

in

a field study of

2004.

Julian Caldecott

unexceptional for gorillas, but the nests were

of the brains of

of

visited

found nests that were built on the ground. This


is

number

had obtained

further surveys, but a

Wrangham

to

The area was

officials

Ammann, who

were said

anomalous

and the body weight estimated from evaluating a

and Bondo,

never to charge them

anatomy but chimp-

cranial

in

chimpanzees {Pan

heaviest

anomalous behavior by the

wildlife

Casts

photograph

1996 by Karl

photographing them; they seemed

gorillas

longest recorded for either gorillas or chimpanzees,

skull there but also collected anecdotal reports of

'gorillas'

crest.

which European hunters

and

team working

videotaping them,

IG.

in

19th century.

in

in

large chimpanzees, one with an unusual sagittal

an area surrounding the

the

swampy

their fur."" Williams revisited the area in

is

its

killed gorillas

skulls

gap

benngei] and the western

inhabited by

is

in

The animals

postcranial anatomy, and to be

midst

troglodytes]. In

towns

known

the

eastern gorilla [Gorilla

is

gorillas.

year, however, a

and pronounced them

DRC

River, there

unusual for

with Shelly Williams succeeded

with Groves,

The unknown ape

is

themselves were not seen.

Flores might not be entirely unconnected with a

are related to

chimpanzee-like, but with a pronounced

with

modern

the thought that a confirmed, small, recent hominin


in

is

sagittal crest.

used

for

laboratory-based

biomedical research.

They are uniquely susceptible

to

human

and serve as a model

hepatitis

for this global

development of conditions related to Alzheimer's

infections

and Parkinson's syndromes, and other forms

health problem. Hepatitis research using chimp-

of

neurological degeneration with age.

Chimpanzees

are the only great apes currently

anzees has
hepatitis

led to vaccines to protect people

B and has also played an important

from
role

27

World Atlas

of Great Apes and their Conservation

in

the development of assays to reduce the risk

of

transmission of the hepatitis C virus through

blood

Other areas

transfusions.

immunodeficiency virus
have proven

to

respiratory viruses.
of

in

although chimpanzees

(HIVl,

is

in

chimpanzees has also become established."

labs for

infants being captured by

w/ild. v^ith

members. Chimpanzees
of

Convention

the

Endangered Species
(CITESl

in

on Appendix

Wild

in

Humane

an

official

the United

and

is

a drive for a Europe-

Society of the United States

ban on the use

permanent

for the

USA, approximately

remain

in

research

Kingdom

several other European

labs.^'

apes

of great
in

the United

relocation of apes

to suitable sanctuaries. In
1

300 chimpanzees now

The National

Institutes of

Health maintain eight National Primate Research

Centers

for studying

nonhuman

primates, of which

two use chimpanzees. The US federal government

Fauna and Flora

spent US$25-30 million on chimpanzee research at

commercial purposes. The

last

23 institutions

such

is

Europe was a controversial decision by

in

2001. Although a large total, this

less than 10 percent of that spent on research

using

monkeys. Fewer invasive procedures are


research involving chimpanzees than

Austria to issue an import permit for 20 chimp-

used

anzees from Sierra Leone

case for other primates,

in

in

ethical

medical and

in

in

Trade

International

of

was banned

biomedical research and testing

the

kill-

1977. thereby banning international trade

in

for primarily

case

Vi(ere listed

on

It

forbidden

is

from research institutions

the mother and any other defensive family

ing

calling for

States,

demand from biomedical


chimpanzees was largely met by imports

Until the late 1970s, the

from the

1997 and

Union countries, and there

genomic analyses

of

resistance on

increasing

is

the use of captive apes

wide ban. The

neurology; drug testing; and

The value

to

other research.

be poor models for HIV research;

genetics;

cognition;

biomedical

of

chimpanzees include the human

on

research

There
grounds

198A,

permits that had been issued prior

on export

in

the

is

probably because of

the apes' cost.^'

to the ban.'-

FURTHER READING
Begun, D.R. 12003) Planet
Brown,

Sutikna,

P.,

T.,

of the apes. Scientific

Morwood,

from the Late Pleistocene

American 289: 7A-83.

M.J., Soejono, R.P., Jatmiko, W.S.E.,


of Flores, Indonesia.

Rokus, A.D. 120041 A new small-bodied hominin

Wature 431: 1055-1061.

Groves, C.P. (2001) Primate Taxonomy. Smithsonian Institution Press, Washington, DC.
Hartwig, W.C. 120021 The Primate Fossil Record. Cambridge University Press. Cambridge, UK.

Janke,

Arnason, U. 12001) Primate divergence times.

A..

All

Apes Great and Small,

London, Moscow,

New York.

vol.

S.,

Marshall, C.R., Witt,

common
Wlldman.

D.E..

ancestor

of

Science

(PNASnOO

at.,

eds,

Monkeys. Apes and Ottier Primates. Jonathan Cape, London.

C, Martin, R.D. 12002) Using the

fossil record to

estimate the age of the

last

extant primates. Nature 416: 726-729.

Uddin, M., Liu,

E. 12004)

to

0.. Soligo,

G.,

Goodman. M.

DNA identity between humans and


Young,

Galdikas, B.M.F., Briggs, N.E., Sheeran, L.K., et

pp. 19-33.

Kavanagh, M. 11983). A Complete Guide


Tavare,

In:

African Apes. Ktuwer Academic/Plenum Publishers, Boston, Dordrecht,

2:

112):

12003) Implications of natural selection

in

shaping 99.4% nonsynonymous

chimpanzees: enlarging genus Homo. Proceedings ofttie National Academy of

7181-7188. http://www.pnas.org/cgi/content/abstract/1232172100v1.

The beast with no name.

New Sc/ent/sn84

12468): 33-35.

ACKNOWLEDGMENTS
Many thanks

to

Peter Andrews (Natural History MuseumI, Colin Groves (Australian National University], Alexander

Harcourt (University

of California, Davis),

David Pilbeam (Harvard University), Christophe Soligo (Natural History

Museum, London), Michael Wilson (Gombe Stream Research


San Diego]

for their valuable

comments on

Center), and David Woodruff (University of California,

the draft of this chapter. Thanks also to Valerie Kapos

research and advice.

AUTHORS
UNEP World Conservation Monitoring Centre
UNEP World Conservation Monitoring Centre
Caldecott. UNEP World Conservation Monitoring Centre

Martin Jenkins,

28

Box

1.1

Box

1.2 Julian

Martin Jenkins,

(UNEP-WCMC)

for

World Atlas

of Great Apes and their Conservation

Great ape
biology
Jane Goodall

Since the
apes

methodological studies

first

natural

their

in

proliferated

in

made

have

differences

of data

has been collected

Africa. Behavioral ecologists

great strides

explaining the species

in

ape social structure and behavior on

in

the basis of differences

and

the

the 1960s, after the Second

World War. a vast amount


from across Asia and

of

suddenly

habitat

in diet,

distribution of food,

from predators and conspecifics. And

risl<s

ongoing studies

flanged males meet

less

dominant males sometimes succeed

consent.

chased away

(obtained noninvasively from fecal samples], and

The apes
range
found

of Asia

of social

and Africa

exhibit a

wide

systems and behaviors. They are

group sizes

in

a flanged

if

from over 100

ranging

male

if

for

bonds between individuals

communities - although

mel

it

bonobos - are

all

gorillas,

smaller groups

in

who

monogamous, maintaining
territories

their relationships

through daily bouts

duet singing and 'dancing'

in

of loud

and

and haunting

the trees. The parents

chimpanzees, and

live in

group contains one or two silverback males,

and a number

families.

Most males also leave

late adolescents,

males

until

mate with males

which they stay

in

to

raise

their natal groups as

sometimes associating with other

they are mature enough to lead a group

own. Then they

may

capture females from

of their

when he or she reaches maturity Siamangs


more social and can be found in larger groups.

other groups, sometimes committing infanticide.

Orangutans, the most arboreal


apes,

live

in

males with

of the great

semisolitude. Fully mature dominant

fully

developed cheek pads, known as

of

their

raise their offspring together, then drive each out

are

of

adult females and young. Young females generally


leave their natal groups, and

and are

(to

of

other groups

pairs

dispersed local

groups

and chimpanzees. Gibbons

in

there

are always together.

the orangutans, and the most social are bonobos


live

in

rather different structures. Gorillas

three to 50 individuals

a few younger, blackback males,

live

Although

many years

highly social, but their groups have

and

apes

arrives.

there are not.

typical

large, the Asian

join

but they will be

would be surprising

(chimpanzees! to lone individuals (orangutans). By

than those of Africa. The most solitary apes are

it,

debate about whether there are permanent

The African apes -

satellite imagery.

mating

large quantities of food are available,

orangutans have been studied

social

DNA

of

When

together with females to exploit

been greatly aided by videotaping, and by new


profiling

in

immature or low-dominance males often

is still

technologies such as genetic

the presence of a receptive

with an ovulating female, but typically without her

ape populations have

of diverse

in

female lethal competition can ensue. Younger or

Chimpanzees and bonobos

live in

multimale,

multifemale groups or communities. These are


fission-fusion

social units, with

ciating in smaller

individuals asso-

temporary subgroups within the

some

chimp-

'flanged' males, live alone in forests that contain the

community range.

home ranges

anzees, particularly females, often travel alone.

of several females.

more than twice the

size of females, advertise their

whereabouts with loud long


visit

These males,

them around the time

calls,

and the females

of ovulation.

When two

When

seasonal

In

populations,

fruits ripen, large noisy

gatherings

congregate to feast together. Chimpanzee and

bonobo males generally remain

for

life

in

the

29

World Atlas

of Great Apes and their Conservation

community

in

which they are born

philopatryl while

known as

(this is

many young females

(like gorilla

females), leave and join other communities before


giving birth. This pattern

mals, but

IS

is

unusual among

many human

typical of

mam-

females

to

in

their group,

and can show

high levels of aggression. Male bonobos are gener-

and there

aggressive,

less

ally

sexual

less

is

dimorphism. Indeed, females sometimes form


alliances

order to dominate males. Conflicts

in

DNA

to

in

gorillas,

defense

in

of

we

share more

our

of

placing us clearly within the ape family

immune

the

of

system, the

and the anatomy

of the blood,

of the

brain and nervous system, are strikingly similar

humans and other


behavior

of

in

apes. This close biological,

evolutionary relationship

makes

the study of the

the other great apes particularly fas-

cinating and important for us, providing insights into

the evolution of

much

our own behavior

of

There can be no question that the apes

semble us

Silverback gorillas are valiant

studies,

The structure

tree.

bonobo society are often resolved through sexual


behavior

some

with chimpanzees than chimpanzees do with

composition

societies.

Male chimpanzees and gorillas are clearly

dominant

According

in

many aspects

of social

re-

behavior They

have distinctive personalities, and show emotions

we

group members, fighting those enemies (including

similar (perhaps identical) to those

human

sadness, fear, and so on. Chimpanzees show

hunters)

who

are not intimidated by their

impressive chest beating, roaring, and fast charge.

Male chimpanzees cooperate

defend their com-

to

munity range, patrolling the boundaries, proclaiming their presence with their loud 'pant-hoot'
distance

call,

and sometimes conducting

lethal

political

behavior (alliance forming and social

communicate with

uage and evidence

of

to attain a goal out

reach of hand, claw, mouth, or beak. Most apes

are capable of tool using and tool making,

modified

object

is

use as

a tool. At

order to

in

one time

this

make

when an

suitable for

it

was thought

to

be the

behavior which differentiated Homo, more than any


other, from the rest of the

has now been seen

and three

of the six

apes. All the

in

animal kingdom. Tool use

the wild

monkeys,

birds,

in

species and subspecies of great

make use

great apes

of

natural

materials to construct their nests, but this


strict tool

nestsl.

use (any more than

bird,

mammal,

not

Bornean orangutans, however, have been

seen holding leaves over their heads


from the

rain.

makers.

for shelter

Sumatran orangutans, chimpanzees,

and bonobos are the most


tool

is

or fish

All the great

prolific tool

ape species, however, are

able to acquire tool-using behaviors

Most fascinating

is

users and

the fact that

in all

in

captivity

areas where

kill,

embracing, patting on

kissing,

the back, swaggering, and so on.

members of other communities - behavior with


many similarities to primitive human warfare.
One behavior that has attracted much attention
an object

and share the

use various leaves for medicinal purposes, and

show

tool use, the use of

mammals

manipulation), hunt

raids into adjoining territories, attacking and killing

is

call joy,

clear comprehension of

have long periods

language

of

of

they

In captivity

human-type langcapabilities.

They

childhood dependence on the

mother, and, certainly

chimpanzees, long-lasting

in

and supportive affectionate bonds between mothers


and

their offspring,

and between

siblings. Like us,

the apes have a dark side to their nature, but they

are also capable of compassion and altruism.

seem

to

We

have inherited both a capacity for violence

and a capacity

for loving

from our shared primate

heritage.

Perhaps the greatest difference between

Homo and

our ape relatives

is

we have
now written

the fact that

developed a sophisticated spoken - and

and electronic - language that enables us

to plan

from the distant past, teach

far into the future, learn

about objects and events not present - even purely

imaginary- and share and discuss ideas. Our highly


evolved

intellect

gives

decisions regarding the


species. Only

us the ability
life

to

and death

make

of entire

we can make the decision to preserve


we work harder to do so, both

the apes. Let us hope

because they are worth

we may

they have been studied, chimpanzees use different

so that

objects for different purposes, and all available

their world,

it

in

their

own

right,

and also

continue to learn from them about

and about our own.

evidence suggests that these traditions can be

described as primitive cultural behaviors, passed

from one generation


vation, imitation,

good

for all

and

ape tool-using

We humans,

30

to the

of

next through obser-

practice. This

is likely to

hold

skills.

course, are also great apes.

Jane Goodalt
Founder, the Jane Goodall Institute

UN Messenger of Peace
www.janegoodait.org

Great ape habitats: tropical moist forests of the Old World

Chapter

Great ape habitats: tropical

moist forests of the Old World


Julian Caldecott and Valerie Kapos

great apes are associated at least to

Allsome
with

types.

Whether

a given tro-

ecosystem supports great ape populations

pical
is

degree with tropical forests, but not

all forest

determined by a mixture

and ecological

factors,

biogeographical

of

combined with patterns

of

habitat conversion and disturbance resulting from

human

activity.

The most important ecological factor

is

the availability of an adequate supply of suitable


food. Here,

relevant that the great apes have

is

it

simple, globular stomachs and lack any special

adaptations to allow fermentative digestion. Other

primate

machs

such

groups,

as

the

monkeys

leaf

have sacculated, fermentative sto-

IColobinael,

(with small 'bag-like'

down

bacteria break

compartments where
has profound

cellulose). This

ecological consequences; these primates are able

from coarse materials such as

to obtain nutrients

mature leaves, which are otherwise


digest and

may

difficult

to

be defended by toxic secondary

metabolites. The diet of the great apes, on the

other hand,

and

to

is

largely restricted to ripe, sugary fruit

other easily digestible plant parts - shoots,

palm hearts, flowerbuds, herbaceous

foliage,

ginger stems, seeds not defended by dangerous

chemicals - and

to the tissues of

vertebrate or

Invertebrate animals.

Body

mammal

the less vulnerable

and the lower

that

digestive

Smaller

the

food

by primates. The bigger any


Iroko Foundation

is,

its

The proportion
leaves)

determines

further

size

supplies needed

it

is

to poisoning

energy demand per unit weight.

of

lower-quality foods

mammal

with

of

the

as

nonfermentative

system can eat increases with

mammals

Isuch

its

size.

same general design

energy. These

Mist rising off the

well onto the great apes: the

rain forest near the

require foods that are

simple rules

map

richer

largest, the- gorillas (around

amounts

in

90-220

herbaceous

kg), eat

con-

village of

Abo Obisu,

smaller

Cross River National

(35-1 00 kg) orangutans favor ripe fruits but tolerate

Park, Nigeria.

siderable

of

foliage;

31

World Atlas

Map

2.1

of Great Apes and their Conservation

Great ape habitats of Sumatra and Borneo

Data sources are provided at the end of

this

chapter

-.-:- '^'-

hi^

I-

'^^^fi

32

<
>

Great ape habitats: tropical moist forests of the Old World

poorer

fare,

such as unripe

and tree bark; and

fruits

the smallest, chimpanzees and bonobos (30-60 kg),

enough

also favor ripe fruit but are agile

may

insects. All of the species

defended

eat young, poorly

ecosystems

of

great ape,

consider

to

is

which they

in

how

the

how

this varies

space and time. The balance and timing

seasonal phases (flowering,

new

of

are thought to

food scarcity
this

of

where

plant

fruiting,

and

fruiting

may

be a serious problem for apes;

if

it

possible to

is

mosaic

move

of forests with

seasonal conditions. Foraging and ranging


in

great apes

is

therefore also strongly

influenced by such patterns, and aspects of social

may

behavior

same

well be indirectly affected by the

to

promote

and

Most animal species native


relatives

[Artocarpusi and

its

(magnoliasi,
relatives,

Sunda continental
islands

of

shelf,

Palawan and Java, and the Malay

Peninsula. The continental shelf

Java Sea, part


of Thailand,

on the Asian or

which also supports the

that are often less than 200


of the

and the

is

covered by seas

deep, including the

South China Sea, the Gulf

Strait of

(dipterocarpsi

the

breadfruit

and the climbing

Borneo are both part

"The

subregion."

of the

West Malesian botanical

ecologies of these islands have

been reviewed by Whitten and colleagues

for

Sumatra," and by MacKinnon and colleagues

for

Borneo

Indonesian

These two

(Kalimantan!. '"

islands both have moist equatorial climates with a

mean annual

rainfall of 2
in

larger size, and

its

mm.

500-5 000

species. Borneo has

Both are

more species

more

of

these are

because
(Table 2.1|.''="'""
else!

of its

on two large

Sumatra (475 000 km'l and


lie

plants,

palms or rattans (Calamoideael.'^ Sumatra and

greater isolation

Borneo (740 000 km^l. These

where

Among

Dipterocarpaceae

families

Magnoliaceae

Biogeography
rain forest

Asia,

groups centered on the Sunda shelf include the

endemic (occurring nowhere

2.11:

Borneo and

to
in

ancestral orangutans originated.

as a result of

Orangutans inhabit lowland

it

local speciation

'^

SOUTHEAST ASIA

(Map

it

species

terrestrial

of

isolation

Sumatra have close

extremely rich

influences.

islands

ages and

ice

biogeography, as at times

region's

within the various islands.''

and

between habitats within

behavior

tended

tree

can be alleviated only

different

successive global

through the Sundaic land masses while limiting

have strong influences on ape

forest

of

phases occur synchronously, seasonal

flushing

to

events. This exerted a strong influence

promoted the dispersion

species

leaves]

Particularly,

distribution.

response

on the

offer the kinds of

live

foods that they are able to eat, and

different

in

warming

dispersion between them, and at other times

key step

the

therefore,

flushing

of

foliage.

To understand the ecology

In

to catch

meat, and are diligent harvesters

vertebrate

years

Malacca, but

is

boun-

Ecology of Sundaic dipterocarp forests

The natural vegetation type


interiors

of

that

Sumatra and

dominates the

Borneo

is

tropical

evergreen rain forest;" this changes with altitude

from lowland mixed dipterocarp forest (below about


700 ml,
to

to hill

200 ml,

about

200

dipterocarp forest (from about 700

lower montane rain forest (from

to
to

500 ml, and eventually

to

upper

500 ml. Each

ded by much deeper water. The shallow modern

montane

seas over the Sunda shelf have come and gone, as

forest type spreads over a wider altitudinal range

the sea level has changed over the last 2 million

on larger mountains, and

Table

2.1

Species richness and endemism


Birds

Island

Number

of native

in

rain forest (above

is

about

more compressed on

Sumatra and Borneo

Mammals

Reptiles

Freshwater

Selected

fish

plant taxa

species

Sumatra

465

194

217

272

820

Borneo

420

210

254

368

900

Sumatra

10

11

11

11

Borneo

48

24

38

33

Percent endemic species

33

World Atlas

of Great Apes and their Conservation

smaller ones. Significant areas

of

low-stature fieath

forest are found tfirougfiout the interior of


in

Borneo

areas of winite sand or otfier very nutrient-poor

soils.

Both Sumatra and Borneo have very large

areas of freshwater

swamp

forests that often

grow

been

Gunung Mulu

The lowland dipterocarp forests are the most

in

have

been

recorded

Borneo, compared

heath forests and fewer than 250


forests."

Root system of a
strangling

fig,

Malaysia.

elevation,

Plant species

although

several hills

in

in

lowland forest but only 12

300 m." The mountains

to
in

in

such

about 850

in

peat-swamp

richness declines with

endemism may

increase;

the northern Sarawak area have

area

bird

The lowland and


so called because

hill

many

at

northern Borneo

of

which 26

to

restricted-range bird species are confined.

species-rich of these forest types; up to 2 300

ecosystems

northern Sarawak, where there

in

comprise an endemic

over deep peat deposits.

species of tree

patterns hold true for birds, for example on

are 171 species


1

The

identified as centers of plant diversity'"

same

'^

'dipterocarp' forests are

of their large trees

belong

to the family

Dipterocarpaceae. The abundance of

these trees

forests

in

is

common

feature of lowland and

hill

Borneo, Java, Sumatra, and the Malay

Peninsula, though Bornean forests typically have

dominance.

the greatest degree of dipterocarp

Their fruiting

which tend

patterns,

among

chronized within and

to

be syn-

species, add to the

ecological influence exerted by their abundance.


Fruiting

is

irregular, resulting in

massive

fruit

crops

Imastings) at unpredictable intervals of two to five

years." This

is

thought

reduce seed predation by

to

overwhelming with food during mast years the


populations of seed-eating animals that are limited
in

abundance by food

other times,

scarcity at

reducing the level of predation inflicted on any one

seed crop.

Water stress during occasional droughts

is

believed to provide the main stimulus for masting

by

dipterocarps.

As many other

tree

taxa

Southeast Asia use the

same environmental cue

prompt flowering, there

is

a tendency in dipterocarp

forests for the foods available to fruit-eating

seed-eating animals

in

to

and

be either superabundant

to

or almost absent at any given time. This helps to

explain

why such animals are collectively rare in


forests compared with other rain
and also why the biomass of fruit-eating

dipterocarp
forests,

primates

inversely

is

related

abundance within forests


similar^''* "
bility

in

The pattern

otherwise

and seed availa-

of fruit

among

depend on such foods. The


to

are

dipterocarp forests favors high

and/or rapid reproduction

them

dipterocarp

to

that

first

mobility

the animals that

adaptation allows

track fruiting activity over wide areas, while

the second allows their populations to respond


swiftly to unpredictable food

supply

Orangutans are fruit-eating animals adapted


to

an environment

in

rather poor supply They

in

which

fruit is

manage

fundamentally
this by being

strongly adapted to arboreality, spending


their time in the trees,

with conditions

in

most

of

and being deeply familiar

the canopy within a large

home

range. Orangutans can therefore track the seasonal

34

Great ape habitats: tropical moist forests of the Old World

changes

in a

patchy ram forest, where the tinning of

peaks can vary with elevation and aspect.

fruiting

They forage within the range


way,

in

unless they happen to

sources

of fruit are to

a typically zigzag

know where

changes mark-

Tropical rain-forest structure

become smaller

edly with altitude, as the trees


In girth

and lower

more densely packed

stature,

in

large

be found.

and with fewer large branches, and the canopy

lowers from 25-40


in

in

lowland forest

15-25

to

lower montane. This structural change alone

would be expected

impose energy costs on

to

very arboreal animal like an orangutan. The total


availability of fruit likely to

be preferred by orang-

utans also declines with altitude, as does


diversity.

floristic

Tree genera that typically comprise major

components

of

orangutan diets progressively drop

out of the canopy composition with

increasing

Nephelium (Sapindaceae), Baccaurea

elevation:

lEuphorbiaceael, Artocarpus (Moraceael, and Aglaia

(Meliaceael

disappear between

dipterocarp forest
[Polygalaceael,

Mangifera (Anacardiaceael, and


disappear between upper

Garcinia IClusiaceael

dipterocarp
1

and upper

hill

700-900 ml; Xanthophyttum

(at

and

oak-laurel

forest

AOO ml' These factors combine

orangutans are generally restricted


less than about 750

(at

200-

to explain

why

to altitudes of

m, except where there are

exceptional concentrations of favored fruit trees."

Trees that provide

orangutans

fruit suitable for

are typically found at higher densities and bear


fruit
in

more continuously

Bornean

iness and

forest,

dynamism

in

in

Sumatran
is

dominance

Sumatra makes space

species that collectively


differences between

forest than

much

patch-

the forests of both islands.

Significantly, the lesser

trees

in

although there

other tree

for

more

fruit

Ian

of dipterocarp

steadily.

Some

Sumatran and Bornean orang-

Single(on/SOCP

large areas of apparently suitable

example

most

in

of

Sumatra, and between the Rajang River

utan behavior have been attributed to the different

Sarawak and the Padas River

patterns of food supply" Figs,

One explanation

such high concentrations

in

in

particular, occur at

some

parts of

Sumatra

may have

that these fruit alone are thought to have enabled

sites

among orangutans

their

greater density and sociability


in

Sumatra than

size,

in

Borneo.' Average home-range

day-range length, and population density

respond

to

differences

in

the

continuity of fruit availability

all

abundance and

between

locations,

in

for this

extirpated

areas

is

some areas

orangutans was discontinuous both


in

in

of

Borneo and

Sumatra. Orangutans are absent from very

those

Indonesia.

of

often contain

other

reluctance to eat orangutans. Examples are


strongly Muslim Aceh Province of

standing
distribution

western Sabah.

that prehistoric hunting

prey.

occur where local people have a strong cultural

between Sumatra and Borneo.


the

forest, habitat of the

Sumatran orangutan,

in

with abundant orangutans

the Batang Ai catchment

deforestation,

in

Primary lowland rain

orangutans; ancient cave

bones along with

Moreover,

for

central

now missing orangutans

seasons, and ecosystem types, both within and

Before

habitat,

the southern two thirds of

hunting

explanation
of

is

in

taboo

in

the

Sumatra and

in

Sarawak, where a longexists.

An

alternative

that orangutans live close to the

edge

an ecological niche that can become unviable

with a slight shift

in

forest composition, for

example

35

World Atlas

of Great Apes and their Conservation

Map

ape habitats

36

2.2 Great

of Africa

Data sources are provided at the end of

this

chapter

Great ape habitats: tropical moist forests of the Old World

in

favor of dipterocarps. According to

patcfilness
lations of

forest

in

argument,

moist forests and rain forests, but their habitats

popu-

span wide altitudinal ranges both within and

in

the

between species, and chimpanzees also use dry

explain

the

forest

orangutans reflects patchiness

may

ecosystem. Each theory

absence

tfiis

distribution of breeding

ttie

and savanna

The African

orangutans from different places.

of

habitats.

rain forests are sustained by the

flow of wet air off the Atlantic Ocean, with winds that

Impacts of
It

is

human

no longer

shed their moisture as they

disturbance

realistic to

describe Sumatran or
to

human

River across the

because the land cover

of both

borders

Bornean ecology without reference


impacts. This

is

seems

islands

be

to

possibly

conversion,

the

In

in

process of rapid

from moist

entirety,

its

forest to plantations, farms, settlements,

and

fire-

maintained grassland. This pessimistic scenario


is

unofficial

oil

palm

illegal],

cellulose used

land

in

Congo Basin

mountainous
the southern

Republic

southern Nigeria (Map

Congo (DRCI

the

of

and savannas intervene, due

ancient interruption

and inland

con-

to the

and prevailing winds - an

known as the Dahomey Gap.

further expanse of rain forest runs

this, a

from Ghana

to

Benin and Togo, drier

2.2). In

figuration of the coast

Beyond

land.

Congo

to the

Democratic

forests

of the

to Guinea, blending further

up the coast

bush savanna, grassland, and

into

ultimately the Sahara Desert.

produce the

These lowland moist forests are known as

and the widespread and

the Guineo-Congolian formations." They are most

of fire

to

means to clear
ever more loaded

species-rich

as a

poverished

is

less

an environment that

with fuel, and increasingly dry as a result of local,


regional,

and cool over

Rwanda and Uganda, from

widespread

poorly planned development of road infrastructure.

The unrestrained use

of

rapid expansion of

mangium,

paper!,

in

in

and

and pulpwood plant-

lElaeis guineensis]

ations (e.g. of Acacia

official

programs,

resettlement

logging (both legal and

use

of land

informed by the recent history

both islands. The latter has involved

rise

The forests stretch from the mouth

and global climate change, has also been

rich

In

in

Central Africa and relatively Im-

West

in

much

Africa. Overall, they are

species than the rain

forests

of

Southeast Asia and South America. Their relatively

low diversity

is

thought

from past climatic

to result

fluctuations that greatly reduced their extent. There

a contributory factor

Given that Borneo and Sumatra are two of

is

good palynological [poUenl and

the most biologically rich islands on Earth, the

of

major vegetation changes

implications of this change for global biodiversity

rain-forest zone of Africa.

are profound, and neither orangutan species could

out repeatedly over geological time, most recently

these circumstances. The

during the last northern hemisphere Ice age, which

only plausible strategy for safeguarding significant

ended approximately 14 000 years ago. These

possibly survive for long

components

of

in

Sumatran and Bornean

biodiversity,

including viable populations of orangutans,

on preserving intact areas

enough

of

conditions. This

is

km^

still

just

1 1

restriction of moist forests

of the drier vegetation

feasible

in

Sumatra

Gunung Leuser

some southern

sand dunes

of the

Kalahari Desert.

forest

000 km' transfrontier forests

of

climatological stresses.

peat

swamps

of

West

West

Many moist

As the climate changed following the

Sarawak
in

types.

parts of the

species disappeared entirely under these

in

in

and

Congo Basin are now growing on what were once

and

11.21,

the

few remaining moist enclaves,

in

forests of the

rich

now

The climate has dried

Indeed, the forests

and the Betung Kerihun National Park

The

evidence

and the expansion

the Lanjak-Entimau Wildlife Sanctuary

Kalimantan.

their species to a

Is

under drought

area (the Leuser Ecosystem, see Box

Borneo within the

changes caused the

fossil

what

lowland forest large

to retain resistance to fire

within the 26 000

depends

in

glacial

maximum, however,

last

the remaining moist

from the few centers

forest species spread out

Those species

were

Kalimantan and Central Kalimantan also have a

where they had

strong potential for biodiversity conservation.

less efficient dispersers tended to remain localized

AFRICA

and endemlsm. The lesser species richness

Biogeography

West African

in

persisted.

that

the refugia, giving rise to centers of diversity

rain forests

is

of the

consistent with both

Together, the African apes occupy a wider range of

the

much

ecosystem types than orangutans.

this

zone during the Pleistocene, and the greater

All of the African

ape species are closely associated with tropical

greater reduction

in

rain-forest area of

prevalence of drier and more seasonal current

37

World Atlas of Great Apes and

their Conservation

enough

days, long

to

purge themselves

of

leaf-

eating Insects. These forests are characteristically


tall,

Emergents

with canopies usually over 30 m.

and sometimes even the main canopy can reach


50-60 m, especially

the wettest coastal zones.

In

Generally these forests are rich

and

poor

relatively

in

tree species

herbaceous and understory

In

except where canopies are disturbed

species,

or otherwise

brol<en.

Aframomum

ginger,

these areas, the giant

In

giganteum, other gingers

members

(ZIngiberaceaeJ, and

Marantaceae

of the

family are notable features of the understory and

A few areas

crucial food plants for apes.

as rich

in

America

woody

widespread, but

some areas
of

Is a

species are relatively

of the tree

concentrations

high

species richness

but, generally, plant

lower Most

little

are nearly

plant species as parts of South

are noted for their

species with

restricted

distributions.

some areas

In

of the

central

Congo

Basin,

lowland rain forests are dominated by one or a

few species

belonging

the

to

legume family

Caesalpinlaceae. Such dominance has major implications for the distribution and foraging ecology of

Gitbertiodendron dewevrei and

animals.

forest

Julbernardia seretii are two of the species that

most commonly succeed

dominating the canopy.

in

These and other dominating Caesalpinlaceae are


noted for producing dry

pattern" and

fruits

having

for

in

mast

fruiting

unpalatable

relatively

leaves. Although these fruits are less attractive to

primates than fleshy


production

and their Irregular

fruits,

poses problems

certain times of year, there

food

of

apes, especially lowland gorillas,


forests

to

nutritious fruits

Inland

advantage

tal<e

when

swamp

move

the

of

they are

scarcity at

evidence that

Is

In

some
these

plentiful

and

season.

and mosaics

forests

into

of

swamp

forests with lowland rain forests also provide key


habitat for apes

make these

Lowland forest

climates, which

chimpanzee

fragmented than

habitat,

in

forests naturally

more

much more broken

canopy
there

rain forests

grow on

well drained soils throughout the region and are

some seasonal

number

of tree

West

Africa.

38

firme forest does and, as a result,

much more growth

story plants.

of

herbaceous under-

The Central African swamp forests

variation

are particularly Important for bonobos, which are

thought

to

enclaves

of

species shed their leaves at par-

They normally remain leafless

of ferra

is

for

more sunlight than the continuous

Although

ticular seasons, they are not synchronized


so.

Central

Iferra firme] forests, but with

Main forest types occupied by African apes

their climates, especially In

in

as the well drained

The Guineo-Congolian lowland

in

especially

tall

penetration of

largely evergreen, despite

Africa,

and uneven canopies. This unevenness allows

Central Africa.

Maiombe, Cabinda,
Angola.

in

They are often as

Africa.^'

in

doing

for only a

few

of

swampy

which

have survived the

swamp

forest.

last

glaciation

in

Elsewhere, local patches

forest are also key habitat for gorillas,

like to

eat herbaceous aquatic plants. The

"

Great ape habitats: tropical moist forests of the Old World

growing

differing pfienology of tfie plant species

swamp

forests

means

in

habitat patcties

tliat tliese

can be very important on a seasonal basis,

in

maintaining continuous food supplies for apes

Throughout the region, forests change


character with increasing elevation. As

decrease

richness, but increase

have greater numbers

stature and species

in

to their

more

epiphytes and a

of

substantial herb layer than lowland forests. The two

main areas

highland forest

of

of

relevance to African

apes are the Cameroon Highlands and the Albertine


Rift

The Cameroon Highlands are volcanic uplands

UOOO km'

western Cameroon

in

and eastern Nigeria. They are

international

of

importance for their endemic birds and amphibians


(Table 2.2|

and also have a high degree

endemism. Due
coast,

and a dense herb-

aceous understory including Gatium ruwenzonense

that are important

to the

of floristic

moister climate near the

in

the diet of mountain gorillas.^"

The bamboo zones

of

these highland systems

number

provide crucial resources to a

animal

of

mountain gorillas and

both

including

species,

eastern lowland gorillas. The bamboo,

Yushania

alpina (synonym: Arundinaria atpina], spreads via

rhizomes and forms a dense canopy, especially


altitudes

bamboo

the

into

season',

rainy

during the

forest

bamboo
when bamboo

season

the

sprouting (September-Novemberl,"

shoots

of

may make up 70-90 percent of their


of the bamboo determines

The persistence

and the abundance

forest structure

it

impedes the establishment

diet.

both

other gorilla

of

bamboo

food plants; the canopy of the


that

at

300-2 600 m. Gorillas move from

of

mixed forest
'short

Highlands.

covering about

revolutum

Hypericum

(Rosaceael,

(Clusiaceael, ericaceous shrubs,

Southeast

in

endemism due

in

in

from other similar areas. They also tend

isolation
to

by Hagenia

grasslands, are characterized

abyssinica

(Rubiaceael, thistles, wild celery, and other herbs

within their habitats.

Asia, they tend to

with

is

so dense

other plant

of

forests occur at lower elevations

montane

(above 500-800 ml at the southwestern end of the

surrounding lowland vegetation

montane

grades

forest

zone

at

Highlands

"

Mammals

Birds

Amphibians

16

20

60

36

30-36

3A

Cameroon Highlands

higher

Albertine

elevations (>2 600 ml on the inland mountains."


Rift

the highlands of Central Africa''

mixed

of

Podocarpus (Podocarpaceael and bamboo

The Albertine

in

savanna. The

is

into

Table 2.2 Endemic species

where the

region than inland (above 2 000 ml,

Highlands

Rift

eastern DRC,

of

Rwanda, Burundi, and western Uganda cover


Thus

around 56 000 km' and include the Itombwe,

species.

Virunga, and Rwenzori Mountains. They are very

habitats besides the

rich floristically

demic

many

bird

and have large numbers

and amphibian species (Table

which are

of

at risk of extinction

rates of deforestation.

The lowland

en-

of

due

to

The

high

distribution of

and transitional or submontane forests that occur


up

to

about

floristically

forests.

Above

600

to

the

in

altitude are closely related

lowland

Guineo-Congolian

They have canopies averaging 30-40


1

500-1 600 m,

which has less than

montane

half as

forest

is

many woody

tall.

found,

species

as the lowland forests and forms a canopy about

15-25

presence
2 100

tall.

These

of coniferous

Podocarpus species. Above

m, bamboo appears

the frequency of
altitude;
'elfin'

forests are notable for the

bamboo

forest

in

the

bamboo stands
is

forests,

and

increases with

interspersed with dwarf or

and subalpine scrub above 2 500

and may form continuous bamboo


elevations above 3 000 m.

forests

may

adequate

to obtain

origin

of

some debate.
bamboo stands

It

This

life

i.e.

cycle

are the subjects of

has been suggested that the

establish only

disturbance, which

monocarpic,

and factors determining the

bamboo stands

may

in

include

response
fire.

of

mass

flowering

at

infrequent

maintain these pure stands as dead

promotes

fire

are

they have only one fruiting season.

intervals of 15+ years, followed by death,


to

to forest

Bamboos

may help
bamboo

that helps to suppress other species.

bamboo stands
become established
during episodes of bamboo

Conversely, the density of the

means

that other trees can

within

them

only

dieback. These episodes

may be

periods of sig-

nificant food scarcity for local gorillas.

Chimpanzees are notable among the African

at

apes for using drier tropical ecosystems as habitat.

The high-altitude open

forests and shrublands, which

bamboo

food supplies throughout the year''^

2.21,

forests

rain

must use several other

gorillas

also be mixed

Particularly

and

Malil

in

and

the far west of their range (Senegal


in

East Africa (United Republic of

39

World Atlas

of Great Apes and their Conservation

Tanzania)

make use

they

and

dry forests

of

woodland mosaics as welt as savanna woodlands.


seasonally

Tfie

dry

forests

West

of

Africa's

supplies. All of the African apes are to a greater or

lesser degree folivorous.

mature leaves

Sudanian belt" are now much fragmented by

at

dense human settlements and agriculture. Their

forests, creates

character

determined by

is

strongly seasonal

annual

rainfall confined

months

eight

almost entirely

Similarly,

to six

or

characteristic

more intense by the

made

is

influence of the hot, dusty

gap environments where herba-

ceous species thrive and provide palatable

of

year The dry season

of the

by the natural treefall dynamics of the

least

mm

distribution of rainfall, with the 900-1 200

but cannot digest the

canopy tree species. Disturbance,

of

some

the fruits of

regenerating forest,

of

such as

Musanga iMoraceae), can make up a major part of


ape diets at some times of year'" For these reasons,

harmattan wind which blows from the Sahara,

lowland gorillas, chimpanzees, and bonobos

and periodic extreme droughts are characteristic

good use

of this zone.

The forests that persist

this area that are

acteristically

protected from

deciduous, and

parts of

in

dominated by two

glandulosum and Guibourtia copatUfera. Along


denser gallery

watercourses there

is

can be evergreen

places.

tvlore

because

it

in

forest,

more

It

also clear, however, that disturbance of

is

humans has

forests by

on apes and that

extensive,

if

only

the woodland

is

impacts

significant negative

species avoid areas where

all

now showing

people are active. There are data

even low-intensity selective logging

in

that

Gabon can

cause a significant long-term depression

important as chimpanzee habitat,


is

which

make

and forest edges as

of disturbed habitats

habitat for foraging.'-"'"-^'

are char-

fire

species of caesalpiniaceous tree, Gilletiodendron

of

chim-

panzee numbers." Given that most African moist


forests

where great apes

are

live

now

allocated to

mosaic and wooded savanna that has been created

logging concessions, this adds poignancy to calls

by extensive forest clearance and grazing

for

both

in

the moister Guinean forest zone and the Sudanian


Isoberlinia

(Caesalpmiaceael woodland.

In

these

systems, dense grass cover and the seasonal

regime impede the regeneration


Similarly

in

fire

of forest trees.

some chimpanzees occupy

East Africa,

areas of savanna that result from the conversion

and degradation

of both

wetter and drier

rainfall

mm."

as low as 850

Where chimpanzees

ways

late
to

make

at

an alarming

consequences

inhabit

open

habitats,

and shelter Their use

of

mosaic varies seasonally,

appears that the food sources

of the

more

characteristically dry zone vegetation, such as the

pods

West

of Isoberlinia in

Africa

and Julbernardia

and Brachystegia (Caesalpiniaceae)

may

in

East Africa,

be most important for chimpanzees at the

driest times of

year

potentially disastrous

ape populations.'

IN

THEIR ECOSYSTEMS

apes play

critical roles in

persistence
their

of

consumption

important dispersers
In

one Ugandan

ponsible for a

determining the nature

same ecosystems.

those

of

forest,

of

many

fruit

they act as

forest tree species.

chimpanzees were

forests

ape habitat

(about A5 percent! of the seeds defecated by fru-

givorous primates. ' fvleanwhile, studies

is

chimpanzees and

for others.'"

gorillas disperse large

numbers

supporting orangutans,

of

increasingly

other forest primates, ^^ and therefore play a major

subject to

viable

seeds over far greater distances than

role in maintaining

better farther from the parent tree. This

the species (see Chapter

all of

persist

those species that regenerate


in

turn

contributes to the diversity and heterogeneity of

131.

to

some

extent

in

disturbed habitats and indeed depend on low-level

ecosystem disturbance

" Both

"

land uses. These pose significant threats to

all

Gabon

in

have shown that gorillas are the sole dispersers for

disturbance, exploitation, and conversion to other

However,

res-

disproportionately large fraction

and high-quality dispersers

Like the Asian

that the eco-

the dominant tree species at Lope National Park,

Impacts of human disturbance

African

is

While depending on their supporting ecosystems,

Through

it

with

rate,

for

THE ROLE OF APES

and woodland

and

companies concerned,

the whole process as 'great ape friendly'

systems that can support apes are shrinking

and

habitat patches within a

be found to collaborate with and regu-

closely the logging

as possible. The general outlook

they are heavily dependent on any available trees


for food

to

more

miombo

woodland formations, including areas with annual

40

foliage.

species that are

for a proportion of their food

the forest.

The great apes also have impacts on

forest

structure and composition through their use of

Great ape habitats: tropical moist forests of the Old World

The edge

of the

Volcanoes National Park


in

Rwanda, showing

cultivation right

up

to

the park boundary, and

pyrethrum growing

in

the foreground.
Elizabeth A-

WiUomson

leaves and

branches as both food and nesting

material. Gorillas

in

particular

petuate the occurrence

may

help to per-

the herbaceous plants

of

tionary time,

however, these ecosystems have

changed as forests and seas have flowed back and


forth over deserts

and grasslands, great

and dried and

swamps

again, and isolated

they favor by causing significant structural dis-

have

ruption to the forest as they feed.""

populations of plants, vertebrates, and inverte-

Apes are also ecologically important as


competitors

and,

in

some

predators.

cases,

Chimpanzees and gorillas occurring

in

the

same

forest area

use very similar resources, but

competition

is

this

modified by the greater quantity of

herbaceous material

the

in

gorillas'

direct contact with

because they avoid

Chimpanzees compete more

and

diet

each other"

directly with

other

frugivorous primates, especially cercopithecine


monkeys,'""'^^ and
sources. There

is

may

drive

them away from food

also considerable evidence that

chimpanzees are important predators of


monl<ey species, especially

Box
illas

4.11,'

and

of

of the red

other small

some

colobus (see

mammals. Both

gor-

and chimpanzees also prey on invertebrates,

and may break open termite mounds as they forage.

filled

filled

brates have regained contact with one another only


to

be divided again later Through

all of this,

within

the constraints of their particular lineage, the apes

have survived as well as possible under prevailing


circumstances in each place, sometimes having
to

move, sometimes dying out before a

mountain

river or

and steadily changing down the

barrier,

generations. The present distribution and success


or otherwise of the apes
live is

the

outcome

in

where they

the forests

of this long history of

adaptation

and movement. Their ecologies are defined by


abilities - to

move

in

their

the trees or on the ground, to

and process their preferred foods and

find

to

tolerate others, to invent and use tools - interacting


in

which

environment.

All of

with their social systems and the ways

foods are distributed


this

complexity

is

in their

rapidly being understood, but at a

when the long history of the


may be coming to an end

CONCLUSIONS

time

Great apes belong to tropical ecosystems; they

forests

shape and are shaped by them. Through evolu-

of

tropical moist
at the

hands

humans.

World Atlas

of Great Apes and their Conservation

FURTHER READING
Chapman,

C.A.,

Lambert,

J.E. (2000) Habitat alteration

Kibale National Park, Uganda.


Furuichi,

Hashimoto, C, Tashiro,

T.,

Uganda: examination

Forest,

American Journal

subcanopy survival

Journal of Tropical Ecology

1 1

Jenkins, M. (19921 Biological diversity.

monodominant and mixed

in

the Kalinzu

forests of the

Ituri

Forest,

443-459.
Sayer, J.A., Harcourt, C.S, Collins, N.M., eds, Tfie Conservation Atlas of

Werger, M.J.A., eds (1989) Ecosystems of the World.

H.,

chimpanzees

Journal of Primatology 22: 929-945.

lUCN and Simon & Schuster, Cambridge, UK.

Tropical Forests: Africa.


Lieth,

In:

primates: case study of

of African

120011 Fruit availability and habitat use by

Y.

of fallback foods. International

Hart, T.B. (19951 Seed, seedling and


Africa.

and the conservation

of Primatology 50: 169-185.

pp. 26-32.

14B.

Tropical

Ram

Forest Ecosystems:

Biogeographical and Ecological Studies. Elsevier, Amsterdam.

MacKinnon,

K.S., Hatta, G.,

PouLsen, J.R., Clark,

C.J.,

Hallm,

H.,

Mangalik, A. (1996)

Ecology of Kalimantan. Periplus, Singapore.

Thie

Smith, T.B. (2001) Seed dispersal by

Cameroon. Journal of Tropical Ecology

17:

a diurnal

primate community

the Dja Reserve,

in

787-808.

Sayer, J.A., Harcourt, C.S., Collins, N.M., eds (1992)

Ttie

Conservation Atlas of Tropical Forests: Africa. lUCN and

Simon & Schuster, Cambridge, UK.


Tutin, C.E.G., Oslisly, R. (19951

Journal of

Human

Homo. Pan and

Evolution 28

WCMC

(19921 Global Biodiversity: Status of

White,

F.

Map

L.J.T., Tutin,

from Gabon.

In:

of Africa.

UNESCO,

C.E.G. (2001)

Weber, W., White,

T.C. (1984) Tropical Rain

Whitten,A.J., Damanik, S.J., Anwar,

Lope

in

central Gabon.

to

Accompany

Hall,

the

London.

UNESCO/AETFAT/UNSO

Paris.

L.J.T.,

Vedder,

New

A.,

gorillas

respond differently

Naughton-Treves,

L.,

to logging: a

eds, African

Ram

cautionary tale

Forest Ecology and

Haven, pp. 449-462.

Forests of the Far East.


J.,

Chapman &

Living Resources.

Why chimpanzees and

Conservation. Yale University Press,

Whitmore,

ttie Eartti's

The Vegetation of Africa: A Descriptive l^emoir

(1983)

Vegetation

White,

Gorilla - coexistence over 60 000 years at

597-602.

(61:

2"' edn.

Clarendon, Oxford.

Hisyam, N.(1984) The Ecology of Sumatra. Qzi\a\\ Mada University Press,

Yogyakarta, Indonesia. Reprinted 120001. Periplus, Singapore.

Whitten, A.J., Whitten,

Wrangham,

R.W.,

J.

(1992) Wild Indonesia.

Chapman,

C.A.,

Chapman,

New

Holland, London.

L.J. (1994)

Seed dispersal by

forest

chimpanzees

in

Uganda. Journal

of Tropical Ecology ^0: 355-368.

MAP DATA SOURCES


Maps

2.1

and

2.2

Data are based on the following sources:

Global Land Cover 2000 database. European Commission, Joint Research Centre, 2003. http://vmw.gvm.jrc.it/glc2000.

Data accessed October 2004.

See species chapters

for great

apes data sources.

ACKNOWLEDGMENTS
Many thanks

to Colin

Groves (Australian National University), David Chivers

Cambridge), and Elizabeth A. Williamson (University


chapter, and also to Marc Languy
Rift

(WWF

of Stirling) for their

Eastern Africa Regional

ecoregion.

AUTHORS
UNEP World Conservation Monitoring Centre
UNEP World Conservation Monitoring Centre

Julian Caldecott,
Valerie Kapos,

i,2

IWildlife

valuable

Programme

Research Group, University

comments on

Officel for

of

the draft of this

information on the Albertine

Chimpanzee and bonobo overview

Chapter 3

Chimpanzee and bonobo


overview
Julian Caldecott

There are two


chimpanzee

species of the genus Pan, the


IP. troglodytes], with four sub-

species, and the bonobo

IP.

paniscus]. They

are of a similar size, with aduU males being 30-61

kg

in

cm

weight and 82-91

head and body length,

In

and females being about 35 percent lighter and


A percent shorten" " Males are therefore rather

more

robustly built

than females. Both species have


black

black faces as adults,

as their legs, and no

arms as

fur,

tor of the

two species may have evolved

country habitat," and


the

Congo Basin

(5.0-1.6

Table

in

is

drier periods of the Pliocene

myal or Pleistocene

1.11,

an open-

in

thought to have colonized

11. 6-0. 01

mya; see

during high-latitude glaclatlons.

The chimpanzee ancestors spread through the


drier forests and woodlands In a great arc from East
David

V\/

A female bonobo with


her infant (Columbus

Zoo and Aquarium).

Liggett Iwww-tlavetiggett co:

long

Diagnostic differences

tail.

between the two species include:"'^

bonobos are born with black

chimp-

faces,

anzees with pink ones;

bonobos have red

lips,

chimpanzees have

brown or black ones;


bonobos have hardly any beard on the
adult

chin,

chimpanzees have white beards;

bonobos are

born

with

prominent side-

whiskers, chimpanzees have none;

bonobo adults
is

retain a

apparent only

In

prominent

tail tuft

that

juvenile chimpanzees;

bonobos have short and very rounded

chimpanzees have longer ones with

skulls,

a lower

forehead and prominent brow ridges;

chimpanzee eyes are comparatively deepset


and close together; and
the bonobo clitoris appears large
that of any other ape,

compared

to that of

Based on differences
it

is

in

and

Is

compared

to

shifted ventrally

chimpanzees."

their mitochondrial

DNA,

thought that the bonobo and chimpanzee

lineages diverged 1.3-3.0 million years ago Imyal,


with the median of the range reported as 1.5

and the

mean

2.1

mya.'

''^'The

common

mya

ances-

World Atlas

of Great Apes and their Conservation

woodlands; they occupy


sea level

West

in

range

Hence they have by

East Africa.

in

most widespread

the

far

and the most cosmopolitan ecology

distribution

any great ape, and show many signs

broad constraints

the

their

of

of

of adaptability

and opportunism. They eat a great variety


within

from

of elevations

Africa to 2 600

of

foods

digestive

system. They cannot cope with a large quantity of

mature leaves, which contain both abundant

fiber

and secondary metabolites such as tannins and

Hence

alkaloids.

dominated by

their diets are

ones when

(especially ripe, sugar-rich

some young

flowers, and seeds, but include


algae,

mushrooms, honey, and

mammals and

in

Over 12 study
observed
of

hunt

to

a year

sites,

at least

chimpanzees have been


32 species

which the most important

monkey [Procolobus

leaves,

a variety of small

As many as 330 food

invertebrates.

types can be eaten

fruits

available),

Box

spp.) (see

of

mammals,

the red colobus

is

A.1|.

They have

also been seen to eat 17 other species of primates,

monkeys [Cercopithecus and

particularly forest

Colobus

spp.l, but also

baboons [Papio

spp.). Flying

squirrels

(Anomaluridae), tree pangolins [Manis

tricuspis],

elephant shrews [Rhynchocyon cernei],

and various duikers [Cephalophus


also reported as prey.

It

community members,

in

occurrence, with a

"

It

between

often shared

is

binges' during which hunting

times.'"

all

particularly in response to

notable that chimpanzees

is

It

are

spp.)

almost always adult males

and the meat

that hunt,

begging.

is

is

seem

to

hunt

an almost

daily

much lower frequency

at

other

hard to explain this pattern solely

is

in

Michael Hulfman

ecological

West

Adult female

Africa,

chimpanzee and her

north of the Congo River Meanwhile, the bonobo

offspring,

Mahale

through North-Central Africa

to

Africa,

ancestors became isolated to the south of this

Congo

Mountains National

in

Park, United Republic

became wetter

after the glacial period,

of Tanzania.

bonobo adapted

its

the heartland of the

the

Okapia

okapi,

to

be

became

swamps, and

the rivers,

effectively

forests that

grew up around them as the rains returned."

ebbs away when

and

may be

rooted

in

would have more


during which the

memory

of recent

Dietary

forests,

deciduous

from

through mosaic wood-

forest,

to

dry

savanna

the easily killed prey has been

flexibility,

variation over a

to forget, until

the

coupled with ecological

huge geographical range within

which seasonality

is

important, can only result

very variable foraging and ranging behavior

animals,

a wide variety of habitats,

all

next time.

in

ECOLOGY AND DISTRIBUTION

lands

'fad'

caught and the chimpanzees start

community

humid evergreen

with a 'craze' or a

it

it

hunting behavior Perhaps this shared enthusiasm

(e.g.

area at drier times, and then

live in

that

animals reinforce each other's

descended from lineages that penetrated the same

Chimpanzees

is

which case

current

biology and behavior to survive.

and the four-toed elephant shrew,

among

common

in

on

hunts and excitement about hunting through further

Petrodromus tetradactylus] also seem

trapped

One speculation

social psychology,
in

terms

physiological

and the

Many other species


johnstoni,

river,

Basin. This area

evidence.

or

chimpanzees,

of

typically of

about 35

nevertheless occupies an area with a

rather limited range of 6-15


of the larger

use an area

km^ and

not all parts

ranges may actually be used. Males

50-200 percent greater than that used

Chimpanzee and bonobo overview

by females, and are

more

likely to

be seen near

boundaries, supporting the scenario

of

females

having small core areas within the defended

range

males.' Chimpanzees are very mobile,

of the

and travel an average

of

chimpanzees reduce
size,

each

day.'

time divided

of its

When

parties.

km

about U

community spends much


foraging

home
Each
into

resources are scarce,


range and party

their daily

Is

somewhat

unsatisfactory

primary environments

of the

in

view

of

the different

two species

(the inner

Congo Basin versus semideciduous woodlands


across Africa], and Implies

need

for

further

research on the ecosystems and biogeography of


the inner

Congo Basin, and the feeding and foraging

strategies

of

bonobos

in

many

locations

and
Chimpanzee

circumstances.

SOCIETY AND PSYCHOLOGY

eat lovtfer-quality food items.'

The main habitat where bonobos have been


studied

primary lowland forest, but they also

is

make use of open woodland savanna,


swamp forest, marsh grassland, and
secondary forests
habitats

if

which

in

dry forest,
disturbed,

possible; they apparently prefer

and edges between them, are

The

social lives of

similar

in

many ways,

others.'" " In both species,

natal

forest there in

chimpanzees and bonobos are


but

deeply different

in

Gombe,

United Republic of
Tanzania! to dry savanna

their

woodland tsuch as

community and migrate from one community

Bating Reserve, Mali).

The Jane GoodaU

young females leave

Institute IJGII

ecosystem types,

variety of

habitat

ranges from humid

spend more time feeding, and more frequently

available. '^

Bonobos

are mainly frugivorous, although their diet also

includes

leaves,

seeds,

flowers,

pith,

mushrooms, and

sprouts,

nuts,

algae. Additional food

sources such as high-quality terrestrial herbaceous


vegetation, earthworms,

termites,

larvae,

ants,

honey, truffles, aquatic plants, invertebrates, and

consume small mammals

occasionally, including

flying squirrels, infant duikers,


is

little

activity

evidence that hunting

bonobos as

for

Bonobos
kill

Bonobos also

have also been reported.

fish

it

and bats, but there


is

monkeys

Interact with

as important an

for

Is

chimpanzees.

at times,

but have not been seen to eat them."

'*

and may

Bonobos

eat less animal protein than chimpanzees, which

may be

related to their greater use of the protein

contained

in

nonreproductlve plant parts," espe-

the stems of aquatic or amphibious herbs and

cially

grasses

in

marshy grasslands.'^

Bonobos

live In

larger than those of

communities that are

than 20-106 individuals


like

is

the range reportedl" but,

chimpanzees, they often forage

parties.

In

smaller

They do spend longer than chimpanzees,

however,

in

large

groups and, since relations

with neighboring communities are far

among chimpanzees,

than

slightly

chimpanzees 150-120 rather

more

relaxed

these groups some-

times Include parties from different communities.


Both species use a mobile, flexible foraging strategy designed to obtain a fruit-rich but otherwise
generallst

diet

of

easily

digestible

and nontoxic

food of great taxonomic diversity, exploiting

many

available

ecosystem types as they

The minor differences


ing

in diet,

as

find useful.

ranging, and forag-

behavior that have been observed cannot

yet be

assessed

for significance. This conclusion

45

World Atlas

of Great Apes and their Conservation

Box

USE OF HUMAN LANGUAGES BY

3.1

Lana joined the research as

CAPTIVE GREAT APES

was two and

believe that the extinction of

apes

preferable to preserving

is

nonhuman great
them forever in

on the grounds that their

captivity,

diminished
great apes

in captivity

nobility

can lead happy

lives, that

value of the preserved genetic material


to

damaged

species. This view

prove

Vi(ill

human psyche would

be very great, and that the

be significantly

the

the

by

loss

these

of

embraces preservation

strate-

gies that create a diversity of niches for great apes

the wild,

include

that

zoos,

refuges,

reserves,

sanctuaries, and even laboratorieslived

in

captive research facility at Georgia State University


in

USA

the

the

work

since 1971, most notably sponsored by

of

Duane Rumbaugh and Sue Savage-

Rumbaugh. This research has explored the mental


and cognitive character

abilities

of great apes, in

the process significantly changing our view of

and how these nonhumans might


modified

landscapes. Two

exist in

Pan

human-

methods have been

human languages

could

Lana demonstrated that she could discrimi-

words and use them grammatically,


create

to

them with

As she progressed, she would sequence

ideas.

novel

utterances

in

planned events that affected her

later starting

response

un-

to

For example.

life.

Lana would request that the research technician

when

her computer vending device

refill

empty

of treats, or

room

was

it

request an item she had seen

that the

computer had no

facility

her Lana exhibited language learning,

to provide to

and her experimental accomplishments were


Equally important to

extraordinary.

lexigram

the

is

developed

keyboard,

her legacy

Duane

by

Rumbaugh, which has served as the primary


communicative interface

for

ape language research

Georgia for the

at Decatur,

This keyboard

is

composed

last several

of three

decades.

panels with

approximately 38^ noniconic arbitrary symbols.

When

the apes depress a key, the word represented

uses sign language; the other, explored here, uses

there

is

graphical symbols that represent words llexigramsl.

IS

to

teach

The following

is

initiatives of the

to great apes:

have participated

in

Research Center

of

the great apes that

the research at the

Language

Georgia State University, and


coexistence with

in

humans.

Sherman and

Lana

project, 1971-1976'''

Lana

is

name

of

language does not

which sought

1970 at the

LANguage Analogue (LANAl

to

develop a computer-based

language training system


the ability of

in

Research Center Her

Primate

in

an

effort to investigate

chimpanzees

to

acquire language.

another before remaining

in

one

to breed.

If

this

LANA

menter and
of

subject.

related

males, but there

is

communication,

relations

in

in

chimp-to-chimp

which they structured their

interactions around statements of planned intent.

Unlike Lana,

Sherman and

Austin could categorize.

with one another, grooming one another frequently


in

hunting,

and sometimes

in

patrolling borders,

killing

neighboring communities, '' and

transfer. Differences

mating with swollen females (see below].

in

the

between males and females.

Among chimpanzees, males

associate closely

in

chimpanzees from

evidence that male

between the two species are most obvious

cuing

The receptive component

language was featured

stalking

chimpanzees also sometimes

human

by focusing on peer

communication rather than that between experi-

and cooperating

closely

and

1973, and Austin, born 197i. using

keyboard. The issue of

would consist

unrelated females and

essence

began working with two young male chimpanzees,

were the only mixing between groups, communities


of

that the

exist outside sociality

was overcome experimentally

female chimpanzee born

derives from the

project,

and the lexigram

Austin research, 1975-1980^

Sherman, born

National

a digital voice

Sue Savage-Rumbaugh argued

the

Yerl<es

spoken by

displayed on a video screen.''

a brief account of the research

Rumbaughs,

the future plans for their lives

humans

one

used

to

chimpanzee. At

believed that only

nate between lexigrams and associate

outside her

Chimpanzees and bonobos have

was

it

use symbols.

is

habitats. Others hold that

in artificial

that time,

when she

a subject

The research was the

interface a keyboard with a

first to

Some

a half years old.

in

guarding and

Among

bonobos. grooming between individuals of the


opposite sex

is

more frequent and occurs

for longer

periods of time than grooming between females or

Chimpanzee and bonobo overview

T^

I^ J liTJ lii

Lexigrams from the 384-word keyboard designed by Duane Rumbaugh for the Lana

pretend, plan, comprehend, and respond to each


other.

complex use

of

language features was an increase

Sherman and Austin

achievements,

not

did

English. Austin died

comprehend spoken

these

Despite

and cooperation.

sociability

in

Sherman's and Austin's more

Attending

in

1998,

Language Research

but the other apes at the

Center have not forgotten him; they

still

make

Oldowan-type rock

project.

with the failure of the chimpanzees

Austin to do likewise.

Sherman and

the basis for the

between the language

difference

by

What was

skills displayed

bonobo Kanzi and the chimpanzees'' Savage-

Rumbaugh had

demonstrated

clearly

reference to him using his lexigram, and they enjoy

and observationally without planned


that

was

This
in

the

first

research

initiative to

language investigations.

caught female
Kanzi- Kanzl

the lab while

It

use bonobos

began with

a nine

month

old baby playing in

Savage-Rumbaugh

tried to

teach his

mother language. Kanzi was not a focus

of the

research because scientists thought him too young


to learn

these

separated

When baby

skills.

from

Kanzi

was

briefly

mother, he began sponta-

his

training;

comprehension precedes production and

drives

language acquisition; and that early ex-

posure

to

language can greatly improve the

competency

level of

attained.

a wild-

named Matata and her adopted son

was

Kanzi

in

language could be acguired spontaneously

that

seeing videotapes of him.

Kanzi research. 1980-1993^'

com-

Kanzi's receptive

tools.

petence for spoken English contrasted dramatically

Panpanzee and Panbanisha research, 1986-1990^


Considering the question
for

questions

to investigate the

a co-rearing study of a
In

of receptive

competence

spoken English, Savage-Rumbaugh proceeded

this

of

species variables

chimpanzee and

Savage-Rumbaugh had hoped

study,

in

bonobo.
to

have Kanzi's mother Matata raise chimpanzee

neously to demonstrate productive competence for

Panpanzee and bonobo Panbanisha

lexigrams and receptive competence for spoken

environments. They were born within three weeks of

English

(something

Matata

had

not

achieved

through direct training].

Kanzi's acguisltion

productive and

competence emerged

receptive

of

following passive observational exposure. Later, as


his

language complex matured, Savage-Rumbaugh

demonstrated that Kanzi's utterances included

grammar,
that

his

learn

males

syntax,

other

only.

and semanticity.

language
skills,

enhanced

It

also

seemed

ability

to

such as the manufacture

of

skill

his

Bonobo males are much more peaceful

in

each other While Matata took good care

identical

of both

babies, she would only allow Panbanisha to nurse.

At that point,

colleagues
they

Savage-Rumbaugh and her human

assumed

the rearing of both babies until

were four years

Based upon

old.

Savage-Rumbaugh determined
the chimpanzees

Sherman and

prehend spoken English

is

this study,

that the failure of

Austin to

com-

not a species-specific
continued overleaf

Female chimpanzees show only infrequent


bonobo social structure

than chimpanzee males, interact less, compete less

social interactions but

for copulation opportunities, are not as territorial,

by contrast, dominated by female coalitions that

are less aggressive with males of other groups, and

Influence

do not hunt other large

mammals. Female bonobos

maintain strong bonds with their sons, which


Itself

In

Increases the frequency of grooming between

males and females."

mating strategies and food allocation.

Female bonobos use


establish

Is,

and

number

maintain

their

of

techniques

bonds,

to

including

sharing food and forming alliances between themselves against males."'

^^

Female

coalitions help to

67

World Atlas

of Great Apes and their Conservation

variable,

Panpanzee and Panbanisha

both

as

floor with chalk;


in

Panbanisha and Kanzi, 1990-present"

Summary

Kanzi lives

in a

bonobo community

includes a 20 ha

wooded

at a facility that

125 ha

forest within a

and theability

of both to participate

musical performances with musicians.

developed receptive competence for English.

An overview

language research with apes during

of

the last 50 years provides strong evidence for their

woodland preserve. The bonobos spend as much

use

time outdoors traveling and communicating as they

as meaningful symbols that refer

to things

do indoors with computers and joysticks. Locations

their qualities (temperature, color,

etc.],

in

the forest are

bonobos know

know

their

named

with lexigrams, and the

this forest as well

own

village.

as

humans might

The bonobos are able

plan where they will go and what they will do

to

when

of

words (manual gestures or graphic patterns!


and

persons

or peers, activities, or as places for foods, rest,


chasing, and so on.

new sentences

with

can use language

Apes can also comprehend

'

fairly

complex structures. They

achieve outcomes that they

to

they get there, and they talk about these plans on

would otherwise not be able

the communication boards.

to

names

formulate

to achieve, for

example

new items based on

for

novel

Kanzi and Panbanisha continue to expand

word combinations. They can use manual signs and

their linguistic world with music, art, writing, tool

graphic symbols to communicate about things that

making, and tool using. Savage-Rumbaugh docu-

are not present; they can learn to

mented on

film

their

breaking

flakes of stone to produce functional

off

Kanzis

ability

to

knapp',

'rock

needs and

to fulfill

language

cutting tools as taught by archeologist Nicholas

their

several years later

manner
Kanzi communicating with Sue Savage-Rumbaugh

come

through the lexigram keyboard. Kanzi's presence at

Its

Great Ape Trust of Iowa

new

in

creatively even

contexts.

If

reared

In

human speech and

understand complex

Language acquisition using lexigrams


optimized
in

apes can

syntax.

the project since infancy has greatly advanced

understanding of apes' capacity for language.

and apply them

skills

that approximates child rearing,

to

for

and games; they can Integrate

specific tools, foods,

Toth; Panbanisha's ability to write lexigrams on the

communicate

one another's requests

if

occurs

it

in

is

the course of social rearing

an environment that

is

language structured.

provides a running vocal narrative to

Ideally, this

the apes as infants, describing what things are.

what

IS

about

should

be

happen, and so on; this narrative

to

symbols that are

show

to

use

of

graphic

function as words.

Results

with

integrated

the

apes can enter the language domain as

that

human

a result of

their capacity for

rearing and Instruction, although

language

than that of humans.

is

much more

limited

great deal remains to be

learned. Future research promises to continue to


blur the boundary between the basic principles of

human and animal


function,

learning, language, symbolic

and complex behaviors.

Duane Rumbaugh and

and are

offset the greater muscularity of males,

maintained partly by a frequent behavior

among

bonobo females: collaborative genital rubbing. This


Is

more

often Initiated by low-ranking females than

by high-ranking ones," and


cile social

bouts

68

of

tension that

aggression or

may regulate and

sometimes arises

when

food

Is

recon-

following

monopolized.

All

ferences

this

of

is

correlated with

sexuality

In

Bill

marked

'building blocks' of sexual physiology.^ In

an age

of

about seven

(in

tfie

same

summary,

bonobosi or 10

chimpanzees], a female begins her


cycle. In

dif-

between chimpanzees and

bonobos. even though the two species use

at

Fields

chimpanzees, during the

first

(In

menstrual

first (follicular)

Chimpanzee and bonobo overview

phase

such a

of

cycle, estrogen

levels rise; this

causes the perineal skin to swell

a very visible

in

that greatly increases her attractiveness to

way

maximum

males. This swelling reaches a


nine days before ovulation and

about three days afterwards. This

maximum

female

(tendency to

proceptivity

swelling

is

the time of

mounting) and
mounting). The

solicit

collapses abruptly after ovulation, as

estrogen levels drop and progesterone levels


thereafter remaining

phase

luteal

follows

as

well

as

attractiveness

(willingness to accept

receptivity

about

sustained until

is

rise,

quiescent throughout the

of the cycle

and the menstruation that

it.

The same process occurs

bonobos, but

in

with the important difference that their swellings

much

last

longer and are

semipermanent.

fact

in

Bonobo swellings vary somewhat during the menstrual cycle in firmness (turgidity) and therefore
attractiveness to males; the swellings peak around

mid-cycle. Female bonobos are continuously receptive

and there

A consequence

that there are far

is

available for sex at any given time

more females
in

evidence that their proceptivity

is little

varies much.^

bonobo community than

chimpanzee

in

one. Since the status of bonobo females

is

not

automatically lower than that of males, as

is

the

situation

for

chimpanzees, and since so many

females are willing and available, male bonobos are


not able to sequester females

each other's access

to

them.

be an

temporary disturbance and may not greatly reduce

both species, a

the forest's carrying capacity for chimpanzees, but

some time
this may

more intense and/or repeated logging causes

for

and

first infant,

important time for her to build social


several communities before settling

relationships

in

down." For

young female bonobo,

last for

up

to six

years while,

in

Sommer

and rarely dispute


In

young female can be sexually active


before she conceives her

Volker

this

phase can

chimpanzees,

it

lasts

mounting disruption
degrading

and sunshine, and increasing

a commercial trade

few months. This implies that a bonobo

there

effects of hunting

whenever her community meets another throughout her subsequent

it

its

up

to drying

winds

vulnerability to

is

those

in

bushmeat, so the

almost inevitably multiply with

oil

locally Increased access also leaves the forest

The most recent estimated

to

in

is

many

173 000-300 000,^ with


but not all areas

in

downward

which com-

parable surveys have been undertaken. The two

West African subspecies are


is

least abundant,

consistent with the greater deforestation

area of distribution. The species

is

which

in

their

threatened by a

that

apes and monkeys

are not scared by the

sudden appearance

of

humans.

open

further hunting and also the fragmentation of the


forest by

mosaic

bound
a

an expanding and eventually coalescing

of

farms and

to find

it

villages.

Chimpanzees are

increasingly hard to survive

in

such

landscape, the more so as the reduced and

fragmented populations come


contact with

other's impacts. Light selective logging causes only

vulnerable to

to

that

are visible from afar so

by land-hungry farmers, leading to

invasion

multiply each

combination of factors that tend

workers

wear yellow hats

ecosystems

CONSERVATION CONCERNS

trends

Gumti National Park,


Nigeria. Field

have similar

extraction can

effects on access, as well as destroying

chimpanzees

field

of logging.

Mining and

life.

total population of wild

fire.

Increased access to the logged-over area along

more

encounter familiar adults

ecosystem,

forest

logging roads encourages hunting, especially where

female

likely to

the

opening

integrity,

its

for only a
is

to

Researchers and

assistants at Gashaka-

into

more frequent

humans and become increasingly


human diseases. Hence, the survival

49

World Atlas

of Great Apes and their Conservation

chimpanzees

of

whole process

ultimately threatened

Is

advancing

of

human use

by the

of tropical

moist forests.

Bonobos are

distributed patchily over a large

common. There

than 100 000 bonobos

are estimated to be fewer

fewer. They are hunted for food

in

some

places.

Hungry

men, and refugees during the


killed

many

the wild, perhaps

in

most places

in

where they occur, although taboos provide


protection

partial

soldiers, militia-

civil

war

certainly

many, but most bonobo populations escaped

More serious was the increased

this direct impact.

some more

1990s, with
to

have

lost

Is

likely

to

hunting from subsistence

the

In

accessible areas

25-75 percent

shift the

to profit,

of

is

likely to bring

expansion

50

d'lvoire,

Republic

Uganda, Lope

in

and Bossou

in

of
In

Guinea.

This fieldwork supplements thousands of person-

years

of

captive

research

on

every aspect

chimpanzee biology (admittedly much

of

it

of

motivated

by the use of chimpanzees as physiological proxies


for

humansi, up

to

and including the Imminent

publication of the chimpanzee's entire

catching up

effort

genome. No

has been directed


species

of this

to
is

fast.

For bonobos,

much more needs

to

be learned

about communication

traffic

In

vocal and symbolic aspects; tool uses and culture;

purpose

of

and

will

keep

other dangers too. Including an

of industrial-scale logging, mining,

and

in

the wild. Including both

and the species' behavioral ecology

in

mosaic

woodland and grassland habitats as well as


forest habitats.

As noted above, further research

needed on the ecosystems and biogeography


inner

Congo

In
is

of the

Basin, and the feeding and foraging

bonobos

many

strategies

Improvements

circumstances. Research using existing captive

new areas

In

access and the spread

of

hunting

(see the Democratic Republic of the

Congo (DRCI country

Democratic Republic of

Cote

United

forest clearance for farming, all with associated

Into

the Congo.

Tai in

the

Budongo

in

bonobo populations under hunting pressure. Peace

bonobo range, Lukuru,

Gabon,

the wild, for example from


in

Tanzania, to KIbale and

their

warfare

of

allow more

bushmeat, however, which may

the

in

Gombe and Mahale

bonobos, although our knowledge

most areas

in

across their range

and trade, and the resulting food shortages

bonobos. Increasing trade with the end

mosaic habitat

researcher-years have been dedicated

comparable research

reported

southern region of

of

the study of wild chimpanzees at study sites

hunting brought about by disruption of agriculture

late

Forest and savanna

Hundreds
to

geographical area of around 3A0 000 km', but are


novtfhere

WHAT WE DO NOT KNOW

profile,

Chapter

161.

of

in

populations, preferably based

with public education,


Jo

is

In

and

DRC and combined


we are to learn

needed

Thompson/Lukuru

locations

If

Wildlife

Research Project

Chimpanzee and bonobo overview

more about language development and


aspects

all

psychically active plants; the origins, role, and signi-

other

ficance of hunting; and the psychological

and communication, as well

of cognition

as the neuroendocrine control

of

sexual behavior

and interindividual relationships among males

research oriented
survival

and females.
As

surer understanding

of

bonobo biology

logical questions

how

new generation

of

on chimpanzees. There are already

chimpanzee biology

way

notably the

flow,

AfricaV

to

of

and the forest

forest

mosaic landscapes can be changed

human

stakeholders
survival.

dissemination of their findings

local

in

to

in

improve

Researchers

should also be on the lookout for ways

to

improve

languages

and other appropriate media. Local acceptance

which "chimpanzee traditions

in

of

of

chimpanzee and bonobo

in

that require further exploration,

from community

the continent of

issues

important and urgent. This

is

management

partnership with

studies

field

many

course,

encouraging and enabling their

the wild

in

the

farming

about differences and similarities

with chimpanzees will surely be raised; this will

then prompt a

to

dynamics

of

would include obtaining a better understanding

is

obtained, a raft of additional behavioral and eco-

ebb and

group existence. For both species,

of

of

the inherent value and interest of the two species of

community, across

Pan

their use of medically or

is

crucial to ensuring their survival.

FURTHER READING
Boesch, C, Hohnnann,

G.,

Ilarchant,

L.,

eds (20021 Behavioural Diversity

Cliimpanzees and Bonobos.

in

Cambridge University Press, Cambridge, UK.


Dixson, A.F. (19981 Primate Sexuality: Comparative Studies of the Prosimians, Ivfonkeys. Apes, and

Human

Beings. Oxford University Press, Oxford.


Fouts, R.S., Fouts, D.H. (19931 Chimpanzees' use of sign language.

Martins

Project. St.

Gagneux,

P.

Griffin,

New

genus Pan: population genetics

(2002) The

In:

Cavalieri, P, Singer,

P.,

eds. The Great

Ape

Genetics 18

[71:

York. pp. 28-';i.


of

an endangered outgroup. Trends

in

327-330.
Goodall,

J.

Chimpanzees

(1986). The

of

Gombe: Patterns

of Behavior. Harvard University Press, Cambridge,

Massachusetts.
Hillix,

W.A.,

Rumbaugh,

New

Plenum,

Kormos,

R.,

(200^1 Animal Bodies,

D.|v|.

Human

Minds: Ape, Dolphin, and Parrot Language

5l<ill5.

York.

Boesch, C, Bakarr,

M.I.,

Butynski, T.M., eds (20031 West African Chimpanzees: Status Survey

and

Conservation Action Plan lUCN/SSC Primate Specialist Group. lUCN, Gland, Switzerland.
Parish, A.R., de Waal, F.B.M. (2000) The other "closest living relative" -

assumptions about females, dominance,

traditional

evolution.

Rumbaugh,

Annals of the

D.M., ed. 119771

Savage-Rumbaugh,
Press,

New

Yorl<

Academy

intra-

how bonobos [Pan paniscus] challenge

and intersexual interactions, and hominid

of Science 907: 97-1 13.

Language Learning by a Chimpanzee: The LANA

E.S. (19861

Project.

Ape Language: From Conditioned Response

to

Academic Press, New

York.

Symbol. Columbia University

New York.

Stanford, C.B.

(19981 The social behavior of

chimpanzees and bonobos: empirical evidence and

shifting

assumptions. Current Anthropology 29: 399-420.

Wrangham,

R.,

Peterson, D. (19971 Demonic Males: Apes and the Origins of

Human

Violence. Bloomsbury,

London.

ACKNOWLEDGMENTS
Many thanks

to Colin

Groves (Australian National University!, Takeshi Furuichi IMeiji-Gakuin University!, and

Hilde Vervaecke and Jeroen Stevens [both University of Antwerp! for their valuable

chapter Thanks also

to

Tim Inskipp and Carmen Lacambra (both UNEP-WCMC!

comments on

to

the draft of this

for research into the literature.

AUTHORS
Julian Caldecott,

Box

3.1

UNEP

V\/orld

Duane Rumbaugh and

Conservation Monitoring Centre


Bill Fields,

Georgia State University

51

World Atlas

of Great Apes and their Conservation

Mike Powles/Still Pictures

52

Chimpanzee [Pan troglodytes]

Chapter A

Chimpanzee
[Pan troglodytes]
Tim Inskipp

chimpanzee [Pan troglodytes Blumen-

The

bach, 17751

known as the

also

is

'robust'

or 'common' chimpanzee to distinguish

whereas the mean weight

was AA

females from Gabon

of 19

kg.

The western chimpanzee

it

is

smaller, with a

from the bonobo [Pan paniscus Schwarz, 1929),

less broad head;

sometimes

Iback of the head], raised brow ridges, and a thicker,

called the 'gracile' chimpanzee, which

much more

has a

limited distribution.

The chimp-

it

has

more rounded, white beard. The weights

anzee has a thickset body, with a short neck and

males were 46.3 and 48.5

broad shoulders, arms longer than

was

tail.

It

legs,

its

and no

The nose

broad and

is

and the hands and fingers

flat

of

two

whereas one female

kg,

only 21.2 kg."'"

The eastern chimpanzee

has a low forehead with prominent brow

ridges and eyes that are deepset and close together

descending occiput

a steeply

is

smaller and

shorter-limbed than the central chimpanzee, with


a

more rounded head, an elongated occiput and


brow

and a

are long, with the outer skin of the middle fingers

straight

thickened. The skin of the face

beard; weights from the United Republic of Tanzania

becoming pinkish brown


fur

is

is

pink at birth,

to black by maturity.

The

long and sparse and mainly black; adults have

and juveniles have

a white beard on the chin


of white hair

remain on

tufts

above the buttocks." " Chimpanzees


fours most of the time, but occa-

all

sionally adopt bipedal postures.

Four subspecies

of

(P.

f.

The

last

troglodytes

verus Schwarz, 1934),

schweinfurthii G'igUoU, 1872), and


IP.

f.

vellerosusGray, 1862).

has also been called the Nigeria chimp-

now thought to be more numerous in Cameroon than in Nigeria. The more neutral
common name Nigeria-Cameroon is used here.
anzee, though

it

is

The central chimpanzee


than the

is

other subspecies,

larger and heavier

with

size

between populations. The mean length


plus body

819
871

in

two sample areas

of

varying

of the

head

Cameroon was

mm and 9U mm for males, and 796 mm and


mm for females." The mean weights of males

were 60 kg

in

whereas three males

from the Democratic Republic

of the

Congo (DRC)

weighed 52.5-61 kg."

The external characters of the NigeriaCameroon chimpanzee are less well known beon genetic characters;"' however, photographs'"

bach, 1799), the western

the Nigeria-Cameroon

52 kg for males and from

22.7 to 45.5 kg for females,'"

cause the subspecies has been recognized mainly

f.

t.

to

but straggly white

Blumen-

IP.

IP,

ranged from 30.3

full

chimpanzee are generally

recognized: the central

the eastern

ridges,

Cameroon and 52

kg

in

Gabon; two

females from Cameroon both weighed 50

kg.

and drawings'^"' indicate that


nent brow ridge and

it

has a more promi-

much smaller

ears.

The

taxonomist Colin Groves has noted that the skull of


this

to

those

and eastern chimpanzees than

to the

subspecies has a closer similarity

of the central

western chimpanzee."
It

has been reported that

could be differentiated from


fifth

subspecies.''

P.

(.

P.

f.

marungensis

schweinfurthii as a

A mitochondrial DNA ImtDNA)

analysis suggested that the lineage of P.

was so
rank.'^'

distinct that the taxon

'

It

schweinfurthii
all

has
is

t.

verus

warranted specific

been argued that as

P.

f.

also diagnosably different, either

three should be recognized as distinct species or

the single species concept should be maintained."

53

World Atlas

of Great Apes and their Conservation

Map ^.1 Chimpanzee

distribution (see country profiles for further detail)

15'N:-

GA^

&H

Species

5'S

Eastern chimpanzee

Central chimpanzee

Nigeria-Cameroon cfiimpanzee

Weslem chimpanzee
Estimated range
^.\S' Eastern chimpanzee
1

^\\'2< Central

ws

ISi'.l'j'

Nigeria-Cameroon chimpanzee

^'.Vj'

Western chimpanzee

250 km

-;-io'&

O'E

54

000

chimpanzee

'

Chimpanzee [Pan troglodytes]

Data sources are provided at the end of this chapter

15E^,

25|"E

NIGER

-^

SUDAN

'"-'l

'VGashaKar,-

'

k'B Gumti

'

'*^

?(FaR) &

<a>

CENTRAL

-s.

>

iv,

tOUBARA
\\'/@C> S
iVv<y CAMEROON J-'X-f^

'
"^

^^?

'Nf^.-i

r~iT--,

.'

AFRICAN REPUBLIC

BFT^;^
^

M_^

/RArihiv

^Koukoua/

.-.-'--^--

fci__'^ .^ai*

KOUIl

'

Congo

V.a_V

/\\

.1

rA

't

i"

^^' ^I

ixJ5*;

vf

T0NG0'Vl|<^u^_4r^

DEMOCRATIC REPUBLIC
OF THE CONGO
;

'

r-,,-T

'V,

,_

['

Like

^''
^^A .
^^tJ*/ Y['f^v->- UNITEDVY)

Mukungu-Vr'^,t5'^t=f5i* ^

'-p.itoniabasi

CABINDA'oa^-

Gombe NP
KASAKATI -

f\

-/

Mahale Mountains

'

NP ^^,

^"~Ccv5^<:

MARUNGU MOUNTAINSri'^ >^>

mM

/^^CV^'.Vi/vJ

\-r

%.

PUBL^H

ANGOLA

'm

vS^'

ri

VI

,^^^

'

r^^Ar

*^

ANGOLA

10"S

10E

20'E

55

World Atlas

of Great Apes and their Conservation

Some

studies based on

mtDNA sequences

questioned

conversely,

between the subspecies


trogtodytes.^''-^^

genetic

the

P.

schweinfurthii ar\d

t.

comparisons

Indeed,

have,

distinction
P.

t.

mtDNA

of

|715N

isolated locality at Toubara

1555'E1,

where

was originally reported in the early 1960s,'^


and was apparently still present after 1983.''
it

Southwards, the subspecies occurs

two areas

in

of

sequences are increasingly leading towards the

Equatorial Guinea, ''' including Monte Alen National

conclusion that there

Park.

is

a very close relationship

between the two chimpanzees


Africa IP.

t.

schweinfurthii ar\6

also between the two


IP.

t.

verus and

P.

f.

of Central

P.

I,

(with

The

P.

of

West

such that

and

t.

IP.

t.

map

occurs

fairly

Congo" and an

troglodytes]

vellerosus as the

and

are

widely

Sanaga

in

(.

It

troglodytes] occurs

extends east

into the

part of the Central African Republic (CAR),


is

largely confined to the

of the

about 4N.

In

the

CAR

it

localities

DRC,

of

north of the

just

in

of the

in

the

Congo

subspecies

is

area."

The western chimpanzee

(P.

t.

verus] occurs

(1258'N

Assirik

1246'W1

southwest

into

1036Wr and

water source |1303'N


Guinea-Bissau."

southeast

in

Mali, north to Djibashin

southern

occurs more or less throughout

It

and much

western

Guinea,'^' Sierra Leone," Liberia,'"'

where

Cote dlvoire,'"' and extends into southwest Ghana,

extreme southwest,

it

in

Dzanga-Ndoki National Park" and several locations


at

also

over a large area southwards and eastwards from

southern Cameroon south

River'"' ^"

It

southern

isolated locality at 3S 16E near

River" The geographic range

Senegal,"
(P.

in

DRC." The southernmost

about 695 000 km'

Mount

fairly

the Kouilou basin

in

the Cabinda province of Angola and

in

extreme west

in

4.1).

DISTRIBUTION
The central chimpanzee

widely

the border with

central, western, eastern

presented here (Map

widespread throughout Gabon'" and the

may

Nigeria-Cameroon subspecies are distinguished


the

is

It

northern part of Congo north of the Equator'''

Africa
it

recognize only two

to

subspecies, the central/eastern

prior namel.^"^

troglodytes],

chimpanzees

vellerosus

one day be appropriate

and the western

t.

and East

has also been found


Juichi

at

an

Yamagiwa

east to about 030'W.'"

of

Burkina Faso, uncon-

In

firmed reports have suggested that chimpanzees

may migrate
country,''"

southwestern part

the

into

of

the

while Butynski"' referred to strong anec-

few chimpanzees were

Eastern chimpanzee,

dotal information

Kahuzi-Biega National

still

Park, Democratic

the village of Douroula. The western chimpanzee

Republic of the Congo.

occurred previously

where

Togo,

1971;"'

and

range

of the

Gambia, where

was

it

it

subspecies

is

has a fragmented range


1

believed to have

about 631 000 km'."'

The eastern chimpanzee

records since

is

recent decades."' The geographic

in

CAR, only one

in

(P.

t.

schweinfurthii]

the north, with few

983: only two localities


locality in

the eastern

in

extreme southwest Sudan,

and scattered

localities east of the

and south

the

to

was appar-

it

of the 19th century;*"

recorded as recently as

last

Benin, where

in

disappeared

in

around the end

ently extirpated
in

that

present along the Volta River near 'the bend' at

Equator

in

Ubangi River

DRC. Between the

in DRC, there are many localities


records since 1983"''"'"^ and there is an

Equator and 5S
with

isolated record

south.

It

in

a small population
Otzi Forest

range

the

extends east

Marungu Mountains

into

was discovered

of the species."'
it

Is

is

the north

Further south

it

occurs

known from the Nyungwe

and possibly from the GIshwati


it

in

In

Reserve, at the extreme northeast of the

Rwanda, where

where

to the

western Uganda,^" where

known from

forest;'"

in

forest

Burundi,

Kibira National Park, the

Mabanda/Nyanza Lake and Mukungu-RukamabasI

56

Chimpanzee [Pan troglodytes]

Landscapes, and Rumonge Forest


Reserve " and in the extreme west of Tanzania"'
Protected

Rukwa

south to the Lwazi River,


3108'EI/'

region (812S

The Nigeria-Cameroon chimpanzee


vellerosus] occurs
highly

southern Nigeria

in

(P.

Hills

Forest Reserve south and east to the southeastern

Niger Delta, and

along the border with

also

Cameroon, from Gashaka Gumti National Park

divisions

Okwangwo and

both the

to

Cross River National

of

the Oban

Country

Bating Iproposed Biosphere Reservel'""

Mall

western

Bossou, near the Nimba f^ountains"^^'

Guinea

western

Budongo Forest Reserve'"

Uganda

eastern

Uganda

eastern

bspecies

Bwindi Impenetrable

is

NP"

NP"

Dzanga-Ndoki

Gashaka Gumti NP
'"

Triangle,

Nouabale-Ndoki NP'"

subspecies

The

also

occurs

in
Ituri

Forest Reserve'"'

western Cameroon, mainly near the border with


particularly

Nigeria,

the

in

Takamanda Forest

Reserve and Korup National Park.''' '" '" This

Kahuzi-Biega

NP"
Reserve"

Kalinzu Forest

Kasakati'"

extends south

population

to

the Sanaga

probably the distribution

is

also occurs

three areas farther inland and north

in

the Sanaga

The geographic range

River."

the Nigeria-Cameroon chimpanzee

U2

encompasses

number

limited

eastern

listed

in

eastern

on

Table A.l.

the establishment of the

in

Uganda

eastern

United Rep. of Ta izania

eastern

NP"

Uganda

eastern

Gabon

central

Mahale Mountains NP""""

United Rep. of Ta izania

eastern

Minkebe NP"''

Gabon

central

Monte AlenNP'""

Equatonal C uinea

NP"

Niokolo-Koba

Assirik,

central

Senegal

western

Guinea

western

Nouabale-Ndoki NP""

Congo

central

Odzala NP'"

Congo

central

Semliki""

Uganda

eastern

Nimba

Mountains'''

research

Early

with

Jane

1967 and leading

Gombe Stream

Studies on western chimpanzees began

TaTNP"''"
"

Cote d'lvoire

western

Tenkere'

Sierra Leone

western

Tongo, Virunga NP'""

DRC

eastern

Ugalla'"

United Rep. of Ta izania

eastern

to

study area

where research continues

the mid-1970s, at Bossou,

chimp-

sites tor

chimpanzees,

Goodalls study beginning

Tanzania,"^

eastern

locations within this broad

of

The main field-study

anzees are

in

DRC

Lope NP''-

Mount

000km'."

distribution.

in

eastern

of

Chimpanzee studies have been focused on

focused

central

DRC
DRC

for the

limit

Nigeria-Cameroon chimpanzee. The subspecies

eastern

Congo

River,
Kibale

about

Nigeria-

possible that they belong to the western

subspecies."

of

central

Nigeria

United Rep. of Ta nzania

Ishasha River^"

which

CAR

C ameroon

Gombe NP"
Goualougo

it

Ch mpanzee

Site

The

Park.^'

western populations are unknown

affinities of the

and

In

chapter

(.

small,

in

fragmented populations from the Oba

southwest

this

The geographic range comprises about

km"'

874 000

Table &.1 Main field-study sites and other locations mentioned

in

Guinea

to

date.

earnest

NP: National Park

Adapted Irom Moore.

and

Tai

J,,

Collier,

M. l]999\ African Ape Study Sites-

http;//weber.ucsd,edu/~jmoore/apesites/ApeSite.html. Updated January 28 1999,

National Park, Cote d'lvoire," and were followed by

work on the central chimpanzee

accessed October 26 200A.

starting in the

1980s, for example at Lope National Park, Gabon.""

Research has continued


ber

of field

understanding

for

many

years at a

num-

allowing detailed demographic

sites,

of the

chimpanzee populations

to

lands

and

deciduous

from

in

East Africa.'"

sea level

in

West

Africa to 2

The

usually important

availability

bution

treatment.

savanna

altitude

has not been the subject

long-term

dry

to
in

be reached. The Nigeria-Cameroon chimpanzee


of a similarly

forest,

woodlands.'^' Their habitats range

but,

in

of

600

year-round surface water


in

some

limiting

chimpanzee

is

distri-

areas, they have developed

techniques for accessing water during dry periods.

BEHAVIOR AND ECOLOGY

At Tongo, for example,

Habitat and diet

well drained volcanic soil and,

Of

all

the great apes,

strongly associated
forests.

They

live in

humid evergreen

chimpanzees are the

with

tropical

lowland

a wide variety of habitats,

forests,

least

moist

from

through mosaic wood-

dig

chimpanzees

up tubers containing

live in forest

when water

is

on

scarce,

water.'"^

Chimpanzees are also very adaptable


face of habitat disturbance.

In

in

the

the Kalinzu Forest

Reserve, for instance, one group occupies logged

57

World Atlas

of Great Apes and their Conservation

Box

CHIMPANZEES AS PREDATORS

4.1

male who either captured the meat or

the

Until

1960s,

are indeed largely

was widely

it

chimpanzees were

believed

fruit

Meat

eaters.

is

that

and they

entirely herbivorous,

consumed

for

only about 3 percent of the time they spend eating,

which

is

less than

nearly

in

human

all

Jane Goodall's pioneering work

documented

chimpanzees

that wild

Today,

Gombe has been

hunting by chimpanzees at
^'^

first

meat and

relish

mammals.

hunt a variety of species of other

documented,'^' -

societies.

Gombe

at

welt

and hunting patterns have

been reported from most other

in

Cote

Uganda,'" and

in

Tai

National

decades

After four

chimpanzees

Gombe,

at

of

research on eastern

great deal

is

known

about their predatory patterns. Chimpanzees

communities comprising 20

to

live in

over 100 animals

that split into smaller parties for short periods of

Such

time.

community

chimpanzees may

of

kill

may be

seven

Gombe chimpanzees

large

kill

numbers
for

of

September,

990. From'

a period of

were observed

late

more than 80 percent

individual

infant

greater chance

mammalian prey An

of

or juvenile colobus stands a


of

being

adult; 75 percent

of

caught than does an


all

colobus

are

killed

68 days, the chimpanzees


colobus monkeys

in

number

of kills, including

those

kill

of the

percent of the

kills.

more

forest

making about 90

hunting,

Females also hunt, though

often they receive a share of

meat from

dominated by Musanga spp. IMoraceael

and with many large

figs [Ficus spp.,

Moraceael,

while another group occupies an unlogged area,


including

forest

dominated

Parinari

by

spp.

(Chrysobalanaceael, mature mixed forest, and


forest."

Some groups

survive

been logged and then almost


agriculture,

where they

travel

in

areas that have

totally

among

converted

of

to

the few small

remaining forest patches and raid crops."

Tomboronkoto region

hill

In

the

southeastern Senegal,

chimpanzees have been found resting and eating


caves during the dry season, perhaps to escape

the high daytime temperatures

in

their

savanna

^iy

human observer
may have been one third greater
time, the chimpanzees may have killed
percent of the

10

colobus

entire

population within their hunting range, a predation


rate that
in

would certainly not have been sustainable

the long term.

The sudden changes

freguency observed at
ecological, social,

immature. Adult and adolescent male chimpanzees do most

early

present,

During this

for

the

in

June through

71

to

hunts. The total

more than

colobus monkey. At Gombe, red colobus account

in

daily

monkeys and other prey

binge seen between 1960 and 1995 occurred


dry season of

however, the red

is,

tend to hunt

unclear For example, the most intense hunting

antelopes each year The most important vertetheir diet

in

to

such binges has always been

resulting from hunts at which no

in

times up

which they would hunt almost

was

brate prey species

at

killed.

animals such as monkeys, wild

and small

of

her early years of research, Jane Goodall"'*

In

and

colobus monkey,

of a single

kill

and eat more than 100 small- and medium-sized


pigs,

is

number

Although most successful hunts result

hunt."
a

The explanation

d'lvoire.^^'^'^

the TaT forests, likewise, there

hunting chimpanzees and the odds of a successful

'binges', during

Park

58

Gombe and

both

strong positive relationship between the

noted that the

Kibale National Park

in

and wolves, cooperation among hunters

lions

Mahale Mountains National Park

Tanzania,

often

other hunting species such as

In

yields greater success rates than hunting alone; in

chimpanzees have been studied: these include


in

from

it

sometimes hunt but hunts are most

female,

social activities.

where

sites in Africa

stole

captor Lone chimpanzees, either male or

the

Gombe seem to

and demographic

in

hunting

be related

Chimpanzees are omnivores, eating


that
to

is

eat

high

in

habitat.''^ In
in

trees

this

was

in

plant foods. Decisions about

meat are based on the

and benefits

of obtaining

prey,

to

factors.
diet

when

nutritional costs

compared

to

the

Bossou, chimpanzees spend more time

the rainy season; a study concluded that

not a response to the vertical distribution of

the food but rather helped

them avoid being

and wet, as they would otherwise be on the


ground away from the breezes

Chimpanzees
an emphasis on
including

of the canopy."'

eat a wide range of foods, with


fruits,

some young

mammals and

flowers, and

seeds, but

leaves and a variety of small

Invertebrates.

As many as 330 food

types Itaxa and plant parts! can be eaten


Diets can vary from

function of what

cold

damp

is

in

a year

area to area, mainly as a

available, but

may

also reflect

Chimpanzee [Pan troglodytes]

essential nutrients that the food provides relative

those available from plants. However, social

to

influences such as party size and composition also

seem
to

A major

hunting behavior

to affect

research on predatory behavior

understand when and why they decide

colobus monkeys rather than forage for

though the hunt risks both

goal of

chimpanzees

in

injury

is

to

hunt

fruits,

even

from colobus

canine teeth and failure to catch anything.


Early studies of this behavior suggested that

meat eating and meat sharing had a strong

social

basis. Hunting was seen as a form

social

display,

in

prowess
the

to

other

members

970s, the

male chimp

which

first

although predatlon

show

his

community.'"

of the

Gombe concluded
chimps was

by

some aspects

based,

tries to

In

systematic study of chimpanzee

ecology at

behavioral

of

nutritionally

hunting behavior were not

of

Craig Stanford

that

A male chimpanzee

at

Gombe

National Park eating

red colobus meat.

well explained by nutritional needs alone. More


recently,

researchers

the

in

chimpanzee research

Mahale Mountains

project

reported

the

that

alpha male there, Ntilogi, used colobus meat for


political gain,

withholding

out to allies.'" At

it

from

rivals

and doling

Gombe, female chimpanzees


meat

consistently receive generous shares of


a

kill

that
after

have more surviving offspring. Indicating a

reproductive benefit tied to

eating. '^^ Other

meat

researchers argue that male bonding


by

it

meat sharing, and

individual

then useful

is

promoted

is

enhancing

in

drawn comparisons between hunting behavior


chimpanzees and
wolves and
to

lions,

be found with

much more apt comparisons are


human hunter-gatherers. In both

humans and chimpanzees, meat

basis whether to hunt. People forage for


also gather plant foods though, as

in

most

of

the organized

opportunistically

when

tor ripe fruit

most

and hunt

they happen to encounter

prey Their meat-sharing patterns are more sys-

personalities all play a role. Future research

tematic and more nepotlstic than behavior seen

in

area should be able to establish further the

underlying motivations and strategies of hunting

In

monkeys, or any other

wild baboons, capuchin

nonhuman

primate.

and sharing. Although most researchers have

local tradition

the techniques used to process

in

food"" and

the

in

There are

and cultural variation."

also differences

medicinal use

plants.'"

of

(^^ovement between foraging sites often takes place

on the ground, but

needed

at

to provide food

The

least

some

and nesting

diet Is usually

from forest trees; the

tree cover

chosen tend
In

is

ripe fruits
to

be those

the form of sugars."'

appears that chimpanzees consume herbs mainly

as a fallback source of carbohydrates


not freely available.'" In the

Craig Stanford

feeding time eating

fruit,

particularly figs, 20 per-

cent eating tree leaves, and a small

amount

time feeding on herbaceous vegetation.'


study
of

In this

figs

of

further

reserve found that at least 15 species

were used, with the

fruits

and young

leaves being eaten."" The foods selected tend to

sites.

dominated by

fruits

with a high calorie content

Is

in

hunting.

hunt; season, group composition, and individual

this

It

meat and

chimpanzees,

there are strong gender biases with males

Chimpanzees forage mainly

male reproductive success.'"

There are many reasons why chimpanzees

only a part of

Is

must be made on a continual

the diet and decisions

societies doing

In

such as

that of social carnivores

when

fruit

Budongo Forest

Reserve, one community spent 65 percent of their

be low

In

be able

monkeys

tannins, although

to

chimpanzees seem

tolerate higher tannin

living In

the

same

forest,

to

levels than do

such as guerezas

{Colobus guereza] or blue and redtail monkeys


[Cercopithecus spp.).""

Some
forms

foods require specific and complex

of processing.

To access the edible part

of a

59

World Atlas

of Great Apes and their Conservation

stream several times

to feed

ex-captive chimpanzees

on algae, '^' and

some

Congo have also been

in

photographed wading."

Lope National Park, insects are an

the

In

important item

in

chimpanzee

diet,

and 31 percent

samples contained insect remains."' The

of fecal

eaten was the weaver ant

species most often

[Oecophytla longinoda]; others included two large


ant species and bees [Apis].

chimpanzees feeding

more time

in

the

In

same

area,

fragments spent

forest

eating leaf petioles, bark, and pith, and

less time feeding on flowers."'

study areas,

up 62 percent

However, as

in all

commonest item, making


food consumed by the Lope

the

fruit is

of the

chimpanzees. They have been recorded eating the


fruit of

U species of plants. When preferred fruits

are scarce, they maintain a relatively high intake


by exploiting small arillate (fleshy, often

of fruit

brightly colored) fruits

The Jane Goodall

Institute IJGII

A young eastern

Saba

chimpanzee feeding on

requires a chimpanzee to remove a thick outer

leaves, United Republic

layer.

of Tanzania.

concluded that

(Apocynaceael

florlda

how

study of

the

example,

for

fruit,

involved are learned

sl<ilis

information

gained

by

food

found

learning hov/ to

imitation

used) or

is

copying

by

learning

(i.e.

it

motor

the

they were recorded to eat 171 different

These were mainly plant materials

Hi

to

species, including 66 species of

Figs were the


in

In

92 percent

most frequently eaten, being

of fecal samples.'"

Mahale Mountains National Park, chimp-

anzees have also been reported eating

soil.

postulated that they do this to obtain

It

is

mineral

behavior involved) without intentional teaching. No

supplements, medicinal chemicals with antacid and

evidence was found

antidiarrheal properties, and to adsorb and detoxify

the

mother or

when

to

suggest that teaching by

imitation by gestural copying

involved.'' Infants

were able

to

were

process whole fruits

alkaloids.'"

is

that

but mastery of

ing others, that eating soil relieves

was

not gained for a

craving for micronutrients, or else

further two years.

^^

those recorded there, have been seen

used by chimpanzees.^'" A variety

may be eaten

when

better

At Bossou, over 200 plant species, 30 percent


of all

The hypothesis

including algae,

termites, ants, and

mammals

of

other items

mushrooms, honey,
such as tree pan-

golins IManis tr/CL/sp/s).'" Fig trees are

most important species

be

to

but,

in

among

times

of

the

food

chimpanzees

have learned, by personal experience or by observ-

they were two years old,

the complex adult technique

In

an unconscious

makes them

feel

they are unwell.

most study populations, chimpanzees have

been observed hunting various prey species, and

meat eating accounts


spent feeding. There

among
fecal

is

considerable variation

populations, however, and the percentage of

samples

tenfold,

for about 3 percent of time

from

to

contain animal remains ranges

0.6

percent at Kasakati to nearly

Gombe

shortage, the parasol tree iMusanga cecropioides,

Moraceae) and the

remains include crowned guenon [Cercopithecus

oil

palm

[Eiaeis

guineensis,

Arecaceae) are commonly used.' Musanga


have been found
other areas,

to

e.g.

fruits

be an important fallback food


the

Kalinzu

Forest

year round. Chimpanzees tend

immersed

in

to

avoid

water but one individual

in

Reserve,

Uganda," presumably because they are available

at

all

being

Mahale

Mountains National Park was noted entering

60

live,

belonging

through observation of their mothers probably


li.e.

Kahuzi-Biega

of

items.

anzees

fruits.

use an object by observing how

forests

National Park, the highest location where chimp-

infants

involves affordance learning

and palm nuts.^"

montane

the

In

percent

at

National

Park.

Identified

pogonias], scaly tailed flying squirrel lldiurus spp.,

Anomaluridae), and

Across 12 study

a duiker
sites,

at

ICepha/ophus sp.)."""
least

mammals have been recorded


most common were primates,

32 species of

as prey, but the


particularly forest

monkeys.'*^

Animals were

killed relatively infrequently by

Chimpanzee [Pan troglodytes]

chimpanzees
16

mammals

as ripe

fruits,

18-30

interact

more

Kahuzi-Biega National Park.

in

month study

one group

there,

killed

In

per year, mainly juvenile or subaduU

Some

Cercop/(/iecus monkeys."

other primates, for

example vervet monkeys ICercopithecus

aethiops],

le.g.

tend

is

opportunities

baboons

IP.

anubis] are occasionally hunted, and

(P.

in

where yellow

the Mahale Mountains National Park,

baboons

olive

cynocephalus] are preyed upon."'

In

and

clumped

that high-energy foods are

can often choose

Gombe, where

intelligent

them

the environment, so that whoever finds

[Papio spp.l are rarely preyed upon. The killing of


at

more

langurs] that eat mature leaves. Part of the

rationale

are often hunted by chimpanzees,'^' but baboons

baboons has been seen only

be

to

- all else being equal - than those

maintenance

who

for

in

first

gains access. This creates

reciprocal

of relationships

and the

altruism

between

individuals,

whether relatedness-dnven or relationship-driven,


or both.

It

is

further argued that large brains are

energetically expensive, so fresh meat, rich

in fats

Budongo Forest Reserve, the commonest


mammalian prey is the guereza [Colobus guereza],

and proteins,

is

an

animal such as a chimpanzee. By

but other prey included blue duiker [Cephatophus

extension to the hominid lineage, an implication

monticola] and an elephant shrew [Rhynchocyon

is

the

Chimpanzees

cernei] captured opportunistically.""

meat

are not carrion eaters, and the scavenging of

from animals

one instance

other predators

killed by
in

Gombe,

a freshly killed

rare. In

is

interest

shown

The predation
by chimpanzees
offers

is

of red

colobus [Procotobus spp.l

discussed

useful example

between

sites.

trees at

Gombe

of

in

detail in

The chimpanzees

Perhaps

chimpanzees only hunt

It

the smaller

in

and tend

to

in

in

that large brain size,

behavior

meat

eating,

and

go together, though as

all

in

reasoning the causality soon becomes

political
all

such

difficult to

untangle.

the

tall

Gombe

response,

opportunistically,

trees

whereas

Ranging behavior

Budongo Forest Reserve, the home range

In

the

of

the Sonso

6.8 km',

community was found

among

ated chimpanzees.""

area

to

cover

may
1

In

contrast,

polygon enclosing the

community

in

to

be about

the smallest reported for habituthe

the Kibale National Park

U.9 km', though

actually have

minimum-

home range

only 7.8

km'

of

was found
of this land

been used. Males used an area

50-200 percent greater than that used by females,

and

were

more

likely

to

be

seen

near

the

Chimpanzees on

patrol,

planned and collaborative

boundaries. This supports the hypothesis that

Kibale National Park,

share meat more actively and

females use small core areas within the defended

Uganda.

chimpanzees adopt

more

i.1.

National Park have to cope with

at TaT National Park.

strategy^'

Box

ecological variation

more aggressive colobus than those

Tai

no

virtually

consuming the carcass.'"

in

particularly desirable food for

bushbuck

[Tragelaphus scriptus] was largely ignored by

chimpanzee group and there was

intelligent

frequently.'*

Males often share meat with other members


of their

group and

this

has been

a fertile

area for

speculation about motives and strategies.

been suggested that the energetic cost

is

has

It

less for

sharing than for defending a carcass, and that


sharing promotes alliances that yield benefits
the form of grooming or support

struggles," or

in

the form

of

in

sexual favors."'

Chimpanzees may also share plant


transfers

are

infants,'" or

usually

in

dominance
These

food.

made from mothers

to

between captive chimpanzees where

some
among mature

food has been supplied by keepers,'*' but

cases

of

sharing vegetable matter

chimpanzees have also been described


wild.'" Sharing of termites has also

mented between mothers and

in

the

been docu-

offspring,"^ and

between adult males.'"


Primatologists

believe

that

species

le.g.

macaques) that consume high-energy foods, such

61

World Atlas

of Great Apes and their Conservation

home range

of the

males." There

is

some

evidence

of

about 35 members, with a range

many

from Budongo Forest Reserve, however, that males

(occasionally as

also spend most of their time within a restricted

social differences

core area.'

populations,

In

comparison with western

which

gorillas,

of

20

over

to

seem

as 1301. There

DO

be

to

between eastern and western


western

with

chimpanzees being

more

generally less violent and

likely to

form stable

generally cover less than 2 l<m per day, chimp-

groups and female-female alliances.'""' There

anzees are more mobile, traveling an average

always a

about
e.g.

li

km

per

day.'"

When

during the dry season

anzees may reduce

of

resources are scarce,

in

their daily

Cote

d'lvoire,

chimp-

range and party

size,

is

dynamic, with

fission-fusion

flexible,

parties forming for short periods and with different

members. This system may make

easier to exploit

it

resources of various sizes, seasons, and locations

spending more time feeding, and feeding more

within the community's

frequently on lower-quality food items."

composition

home

range.'" The size and

of parties is influenced by the threat of

predators, including people; the presence of other

mammal

Social behavior
It

is

hard

to

describe any aspect of chimpanzees'

behavior and ecology

Few

of their

tal<ing

only

one factor

personality, history,

influences

in

in

isolation

day-to-day decisions
into

from any other

seem

to

account;

made

individual

and relationships are

the expression of behavior

be

all

crucial

in different

contexts. The options for behavior are also very

wide,

since

more complex behavior has been

animal.

displayed

list

of the

at

by the Mahale chimpanzees included

resting,

Ta'i

National

Park, Cote d'lvoire.

when assessing

considered

food

to several

the

that both

should

food

of

impact

be
food

of

as a factor

in

determining such patterns

declined with Increasing food abundance.

The

presence

'swollen'

of

females,

with

prominent perineal swellings around the anus and


vulva, ^'

has more effect on party size than does food

can build

common

eastern chimpanzees

supply on grouping patterns.'" The Importance of

were patterns also commonly seen

humans and

abundance

and

dispersal

availability.'

in

of

Budongo Forest Reserve suggested

over 500 descriptive terms, of which more than 200

bonobos, and about 50 were

Western chimpanzees

behavioral patterns

One study

food abundance.

reported for this species than for any other non-

human

species; and the availability and distri-

bution of water and nesting sites,'""" as well as

., 100.275

up when

attracted to a

|_a|-gg

parties of

chimpanzees

swollen adult female

is

party consisting of a top-ranking

study populations of chimpanzees but not observed

male, often accompanied by large numbers of adult

bonobos.'" For more on behavioral differences

and adolescent males, with other males joining

in

between chimpanzee populations, see Box

Ika Herbinger/Wild

the group to be with her'" Meanwhile,

4.3.

A chimpanzee community has an average

size

Chimpanzee Foundation IWCF)

in

small

communities, single high-ranking males may

try

sequester and guard swollen females.

the

to

unusually

In

community at Ngogo, KIbale


number of males Is very

large

National Park, where the

high, pairs or trios of top-ranking

cooperated

to prevent

males sometimes

swollen females from mating

with other males and tolerated each other's mating


activities.

^'^

From these observations,

it

seems

that

such females are apparently so attractive that the

male response

to

them can

other considerations
requires
At

some
the

In

easily

overwhelm

their decision

making;

all

this

explanation.

age

of

about

10

years,

chimpanzee begins her menstrual

female

cycle,

with a

periodicity of about 35 days.'^'^' '' During the first


Ifollicularl

phase

of

the cycle, estrogen

by the developing Graafian

follicle

Is

secreted

- the capsule that

protects a developing egg. This causes the perineal


I'sexual'l

skin to swell

in

a very visible way.

As

estrogen output peaks just before ovulation, so does


the swelling; this

62

is

accompanied by changes

in

Chimpanzee [Pan troglodytes]

behavior and
as

relations with adult males, as well

in

her attractiveness to males. Under the further

in

influence of the ovarian and adrenal androgens,

becomes

the female

preceptive and actively seeks

sexual contact with males, behavior that reaches a


at or just before ovulation

peak
is

when

the swelling

She also becomes receptive

greatest.

mission and the maintenance

of

posture until intravaginal ejaculation

mid-cycle surge

This

preparedness

most

in

estrogens.

the ovarian

is

monkeys], however,

it

achieved.

sexual motivation and

in

conventionally called

is

mammals, and

to intro-

the copulatory

estrus'

in

driven principally by

is

it

In

apes land Old World

is

under more complex

neuroendocrine and social control, so the term


estrus

is

not wholly appropriate.'"

After ovulation, estrogen levels drop sharply

and the corpus luteum develops from the


and starts

to secrete

the swelling

progesterone.

In

The Jane Goodall

response,

and remains quiescent

collapses,

and the

throughout the luteal phase

of the cycle

menstruation that follows

it.

A young female can

have these regular bouts

of swelling

first infant,

her

and

this

may

communities before

expression of a variety

All this leads to the

usually found with

social forms.'" Ivlales are

mixed

in

range atone or

in

are,

however, records from 1976

groups

whereas females

parties,

often

small parties with other females.

mutual grooming, but social interactions between

seems

that

males choose

to associate with

It

partners

on a tactical basis, rather than randomly grouping


or independently selecting the

which

forage.'" There

to

is

same

little

locations

in

evidence that

the large

members

of

boundary

levels of aggression

between individuals

of differ-

ent subgroups.'"
Patrols along the boundaries of the range of
a

community are carried out by groups

may

of

males,

lead to lethal attacks on

communities, often targeting their

males or young."'"' These combative patrols may

community by extending

benefit the

protecting

their range,

other community members, and

more females

corporating

into

these advantages apparently outweigh the risks


involved

in

carrying out the patrols and attacking

conspecifics.

Within a chimpanzee community,

see aggressive behavior that

is

it

is

rare to

intense enough to

community member There are

cause the death

alliance formation. Juveniles and adolescents tend

records from Tanzania, where an alpha male

males begin

mothers. As they get older,

to associate

more with adult males,

killed,'

was

mothers

males

in

early adolescence, before transferring to

80 person-years

in

later adolescence.'"' In over

of field observation of well

known

of a

violently
in

the

he behaved
allies

was

and from Uganda, where the victim was a

young adult male."

while females often continue to associate with their

other communities

in-

the community;

with varying degrees of cooperation, coalition, and

to associate with their

two

which tended

were low

kinship strongly influences male relationships.""


Social interactions are frequently complicated,

in

in

neighboring

nonswoUen

chimpanzees.

natal

to associate in

together and participate

research suggests that

females prefer each other as party members.^'"

of disap-

their

patrols."' Despite this clustering, there

to stay close

Ivlore specific

associating with each other, whereas

particularly

represent emigration

Kibale National Park, 35 males

In

usually adults, and

for

may

Ngogo community were seen

marked contrast with

males show no particular preference

1997

is

important between male

rather than mortality"'

|in

females are infrequent

adult

Bossou, which

at

to

immature males from

of

Grooming

There

to transfer out of his natal group.'"

pearances

Males associate closely with each other, including

the bonobol."

Tanzania, no male chimpanzee has

in

been seen

separate subgroups, the

down.'"

females

individuals

be an important time for

to try out life in several

settling

of

and sexual

months before she conceives her

proceptivity for

UGH

Institute

follicle

In

Tanzania, a young adult male

attacked and ostracized by other

same community,
in

ways

apparently because

that the alpha

male and

his

found provocative.'"

63

World Atlas

of Great Apes and their Conservation

Box

^.2

CHIMPANZEE VISION

prompted the suggestion


to

During primate evolution, vision has become

most

highly developed sense.

chimpanzee

life

and

It

is

such

of forest leaves.'"

Trichromatic species

are indeed better at this task than dichromatic

activities

ones."'^" Trichromacy also benefits folivory," and

of tools, feeding, recognition

of individual conspecifics,

background

of

an integral part

culture, allowing

as the sophisticated use

ttie

that trichromacy evolved

allow identification of ripe fruits against a dappled

and communication.

would have promoted the selective

evolution

Its

exploitation of

young leaves (which are often red

in

tropical forests) as a food resource."

Vision and primate evolution

The

first

primates probably evolved from insecti-

mammals more

vorous

than 65 and perhaps as

Visual cues and social interactions

Frugivorous primates have relatively large brains

long as 80 million years ago Imyal. The theories

with

surrounding the subsequent adaptive radiation

of

primates include the development

of

enhanced color

of

vision as a key step, with various explanations for

The presumed arboreal

it.

primates led

lifestyle of early

more neurons

in

the parvocellular system, one

the pathways for processing visual information.'

This system primarily analyzes fine detail and color,

supporting selection for the

early researchers to suggest that this environment

the parvocellular system

promoted an increased reliance on visual and

mates with larger group

tactile

senses.'^^'^" Furthermore, orbital convergence

and

IS

critical

with nails rather than claws, could be particular

facial

adaptations for a visual predator"

to

canopy

at night, increased visual acuity

the forest

would allow

also observed

is

in

social

in

pri-

system

sizes.' This visual

therefore likely to be used by chimpanzees

stereoscopic vision, together with grasping hands

In

detect fruits

ability to

based on specific visual cues. The same trend

in

such as interpreting

interactions,

expressions and gaze direction. The

recognize other individuals

development and evolution

of

crucial

is

mammalian

ability
in

the

social

discrimination between potential food items. Precise

systems. Chimpanzees also have the remarkable

eye-hand coordination would also

ability to identify

operating

in

the dark. Locomotion

been an important force selecting


vision

locate

in

animals

may

also have

for stereoscopic

arboreal animals that would

branches or trunks

between them.^* Today,


IS

aid

precisely,

this

in

enhanced

need

order

to

to leap

visual acuity

used by chimpanzees when manipulating

tools.

known

mother-son relationships

in

un-

individuals, using only visual cues. Vision

mechanism

therefore an important

of kin

is

recog-

based on outward appearance, independent

nition

of previous experience."" This ability

is

very impor-

tant to

chimpanzees, where related individuals may

spend

some time apart and where political


may be formed on the basis of familial

alliances

Color vision

relationships.

Primates have excellent color vision compared with

most mammals. Chimpanzees,

other

communication, both

apes and Old World monkeys, have color

vision

sions and

based on three different types


in

of

cone photopigment

the retina of the eye.''" Each of these photo-

pigments absorbs
and

a different

this type of color vision

Trichromatic

primates

discrimination

in

is

wavelength

have

good

particularly

to

primates with only two

types of photopigment Idichromacyl.

Infanticide

of light,

known astrichromacy

the red-green part of the spec-

trum when compared

has been reported

populations across Africa, but

eastern chimpanzees.

It

is

in

This

chimpanzee

Gombe

in

in

the

National
"'
Park,"' at Mahale Mountains National Park,'^'- '"
at

'language'.

in

chimpanzee

the gestural basis of chimpanzee

The importance

communication
that

in

determining facial expres-

in

is

of vision in

chimpanzee

supported by the observation

chimpanzees are

readily

communicate hundreds
researchers even believe

able to learn and

human 'signs'. Some


that human language has
of

evolved from a gestural origin, which would have


relied heavily

on vision."

has

most frequent

has been recorded

Budongo Forest Reserve,'"'""

64

Vision has an important role

like

that of

Alison Surridge

and

Kibale

at

usually

National Park.'" '" Infanticide

carried

out

by

males,

and

is

has been

explained as a strategy to bring the mother back


into a

condition of

earlier than

fertility

and sexual

receptivity

would otherwise be the case, poten-

Chimpanzee {Pan troglodytes]

increasing the

tially

chance

killer's

of fathering

have only three or four offspring during their

to

her next offspring.'"' The dead infants are often

lifetime. In the Tai National Park,

subsequently eaten, with the meat being shared

invested about two years

as usual."' ^" This suggests that nutrition

caring

land

sons, whereas

for

about

the enhanced status that goes with control of a

invested

meat resourcel may be another motivation

daughters."

for

infanticide." Observations of infanticide occurring

within a Mahale

community showed

that victims

were always unweaned males whose mothers had


mainly mated with older adolescent or immature
males." Kidnapping

of infants

by males has also

been observed, and may result


starvation

unweaned

of

suggested that

rather than an interest

in

can be an important cause

thought

means

in

it

a statistical

the norm, and

is

important

chimpanzee

in

major

coalitions

that
in

it

is

role in servicing relation-

improve status.^

can

clusters varying from two

23 individuals, with adult males grooming tor

longest

small clusters, and adult females

in

grooming

clusters of five or more.'"

in

has been noted

of social scratching

with grooming

Park"^ and
not at

at

in

association

the Mahale Mountains National

Ngogo, Kibale National Park,'" but


National Park, Kanyawara (Uganda],

Gombe

or anywhere

in

A custom

in

West

29 years

over

Chimpanzees from communities

that have had

of

infant

daughters, and

more

was recorded among 135


study

of

for

births

Mahale Mountains

in

National Park.'"

The demography
has been studied

cause

death

of

Mahale chimpanzees

of the

The major

in detail.'"

was disease

identified

(48 percent), followed

by senescence (24 percent), and within-species

aggression (16 percent). Half

history include the first

an average

groups

at

of

1 1

at 13 years.

The fecundity

of

female

maximal swelling
to

at

other

first

time

females was highest

old, with a birth rate of 0.2

similar study at Bossou from

2001 found that the average age of giving

birth for the first

than

chimpanzees
typical

years, and giving birth for the

per female per year


to

in a

about 10 years, emigration

between 20 and 35 years

1975

of all

Landmarks

died before weaning.

in all

time

(at

about

years)

was lower

other wild chimpanzee populations, and

that the infant

and juvenile survival rate was the

highest. This suggested that the lifetime repro-

ductive success

was

likely to

be higher

than at any other long-term study

Members

Africa.

caring

young.'" Twins are rare: a

of

single twin birth

life

among males, where

especially

Chimpanzees groom
to

death. Although
death

weaned young.

to play a

ships and

of

by no

is

Mutual grooming
societies,

its

faster-maturing

production

rapid

has been

It

possession' or plaything

interest in the infant as a

rarely affects

the deaths by

sometimes motivated by an

this is

sense,^" infanticide

in

individuals.

in

National Park, high-ranking

females have significantly higher rates


survival,

in

subordinate females

months more

11

Gombe

In

dominant females

more than the average

of

at

Bossou

site."''

with one another, with extragroup paternity being a

frequent contact with

minority event.

from those that

humans behave differently


encounter humans for the first

Andrew Fowler

time. Encounters

in

This Nigeria-Cameroon

each community breed mostly

study

in

the Tai National Park,

infant will

dependent on
until

it

is five

its

remain

mother

years old.

the Goualougo Triangle (Congo)

with chimpanzees that lacked prior experience with

humans were
of

characterized by a high frequency

curious responses.'"

Where chimpanzees have

had more contact with people, the survivors tend


to

be very

much more wary

of

or aggressive

towards humans.

Development and reproduction


Wild chimpanzees have a very low reproductive
rate.

of

Females reach sexual maturity

age and

thereafter

mothers
a

life

mean
is

typically

Infants

give

birth

r^m

-^

**^

10-13 years

every six years

are very dependent on

for the first five years.

expectancy

at

'^'9-^^-^

their

They generally have

at birth of less

than 15 years; the

adult lifespan, after reaching sexual maturity,

about 15 years.'" Females are therefore

likely

65

World Atlas

of Great Apes and their Conservation

Box

4.3

CHIMPANZEE CULTURES

parasites, social customs, and courtship gambits.

This complexity contrasted strikingly with earlier

Midway through the 20th


about

nothing

to

Subsequent decades
term

sites

chimpanzee

possible to put together a

it

comprehensive assessment

much

across Africa,
for

human

of behavioral variation

as anthropologists have done

societies.

chimpanzees

next

behavior.

of field study at multiple long-

made

have

we knew

century,

wild

The resulting picture shows

have a rich cultural complexity that

to

was unsuspected

in

pathway

this

discovery

of

included attempts to identify differences

feeding

in

Gombe and Mahale

habits between the

sites

in

Tanzania' and the identification of a social custom,


the

'grooming

handclasp',

which

Mahale but not Gombe. '^^ As more long-term

field

studies progressively yielded greater knowledge of


local habits, researchers

began

draw up com-

to

parative tables that suggested a growing

behavioral differences right across

which

traditions,

Moreover, each community was found

to

that,

these traditions so

of

know enough about an

we can now

assign

to

to its

it

of Its cultural profile

list

"

Africa."'-

of
'^^

In

were

where long-term

field

first

time, to create a

to

more comprehensive under-

phase,

list

In

suggested patterns

suspected might represent local


ating a

the

of

first

phase,

behavior they

traditions,

65 'candidate behaviors'.

of

the sites

pool their data for the

standing of local traditions.'^"


site directors

all

studies on chimpanzees were

being undertaken agreed

In

gener-

the second

each behavior was coded by the core

researchers

at

(performed by

each
all

as either cusfomafy there

site

phenomenon

human

of

among

able-bodied individuals

several individuals,

some degree
either witti

of

of at least

consistent with

social transmission), or absent,

an apparent ecological explanation [such

as the absence

of

appropriate raw materials! or

no such explanation. Based on

w/f/i

this last piece of

information, putative traditions or 'cultural variants'

were defined as those behavioral patterns


either

customary or habitual

chimpanzee community,
logical explanation at

yet

in

The 39
forms

were
one

another Genetic explanations

of

notably

criteria,

behavioral variants.

cultural variants

of tool use,

that

least

absent without eco-

were also excluded using various


geographic proximity

at

one

is

of

we

identify with

culture, our inevitable

reference point for such comparisons."'


This systematic two-phase procedure

being applied

additional

is

now

the second Collaborative Chimp-

in

anzee Cultures Project, which

first

is

investigating

Areas

behavioral variants.

omitted from the

explicitly

such as

study,

styles

of

and carnivory. are being included. The

hunting

to

orangutans

in

effort that

use and social customs. For both orangutans and


chimpanzees, there

identified

included

techniques for dealing with ecto-

is

evidence that the variants are

socially transmitted. First, the similarity of overall

behavioral profile

correlated with

is

geographic

proximity, indicating that behaviors are transmitted

from the location

at

which they

first

communities with higher indices

appear Second,

of social proximity

have larger cultural repertoires. Where apes spend

more time
learn

are

together, they have greater opportunity to

new behaviors from one

more

likely to

another, so behaviors

spread throughout a group.

These discoveries are

one age-sex category], habitual (occurring repeatedly

expect

identified 19 cultural variants, including both tool

inevitably limited.

the late 1990s, the directors of

we

to talk of 'culture'.

precisely this multiplicity that

IS

the

makes sense

it

procedure has also been applied

site,

we

community on the basis

alone (see table], as

an equivalent international collaborative

have been published from each

if

do for people. Indeed, the fact that each com-

the reasons that


It

exhibit

chimpanzee,

individual wild

happened

to

typically

behavioral variation.

single

However, as these were based only on the data that

the conclusions

66

animal

around 10 or more

present at

is

of

recognized just a

munity expresses these multiple traditions

before the fieldwork began.

Early steps

reports

significant

for

both

anthropology and conservation. From an anthropological perspective, the

understand the roots


cultural capacity

The

ancestor of about 15

in

new

picture

helps us

our own extraordinary

common human/great ape


mya was

something

bited cultures

seen

of

like

likely to

have exhi-

the simpler forms

chimpanzees and orangutans today From

the consen/ation perspective, the tragedy


not only do

we

risk losing several

is

that

subspecies

of

chimpanzee, but also and more imminently we


risk losing their

richness

unique subcultures. As this very

becomes more widely known, perhaps

greater consen/ation effort will be mobilized.

Andrew Whiten

Chimpanzee [Pan troglodytes]

cultural variation across long-term study sites

Chimpanzee

most long-term study

Distribution across the six

being customary or habitual


explanation

at least

in

in at least

sites of 38" behavior patterns that

one community, and absent

Guinea
Pestle pound Imasfi palm crown

Rain dance [slow display

stone

inut-hammer. wood

hammer

on wood

hammer

on wood
on stone

anvil)
anvil]

anvil!
anvil)
etc.)

other [food-pound onto other

(e

Termite fish-M

(J

[termite-fish using leaf midrib)

Termite fish-S Itermite-fish using non-leaf matenalsl


Fluid dip (use

of

probe

to extract fluids)

Ant dip [one handed

dip stick on ants)

Ant fish [probe used

to extract ants)

Ant Wipe (manually wipe ants

Aimed throw

Index
Fly

hit

Leaf clip

to fan

mouth

[leaf

Leaf napkin

[np parts

of at

one age-sex class

le.g.

adult malesi

Habitual: has
in

several individuals,

consistent with

degree

some

of social

transmission

o
o

Present: neither

customary nor habitual


but clearly identified

Not possible:

absence can be explained


by local ecological

marrow

out)

o
o

o
o
o

o
o
o
o

Absent: not recorded

with no apparent
ecological explanation

Not known: not

recorded, perhaps

with mouthi

because- of inadequacy of

relevant observational

opportunities

dabbed on wound, examined)

off

[pull

Seat vegetation

stems

leaf!

O
o

noisily to attract attention)

(large leaves as seat)

(tickle self

using objects)

(slap branch, for attention)

(clasp

o
o

arms overhead, groom)

Knuckle knock (knock

to attract attention)

(strike forcefully with stick)

stem, as threat)

[inspect ecto-parasite on leaf)

Pull through

Club

(S

least

most able-

members

features

leaf with fingers)

Leaf squash [squash ecto-parasite on

Branch slap

or

[leaves used to clean body)

Leaf inspect

Hand clasp

o
o

o
o
o

flie

off leaf,

Customary: occurs
in all

occurred repeatedly

items underfoot, courtship)

Leaf strip (np leaves

Self tickle

o
o
o
o
o

o
o
o
o

[intense 'grooming' of leaves)

Leaf clip fingers [np single


Leaf dab

wandl

pick bone

to

used

[leafy slick

groom

Tanzania
<s

s
s

[disable bees, flick with probel

Shrub bend [squash


Leaf

off

o
o

[squash ecto-parasite on arm)

whisk

Mahale Gombe

(throw object directionally)

pick (probe used

Bee probe

(S>

enlarge entrance)

to

stone))

Expel/stir Istick expels or stirs mseclsl

Marrow

Q
O
o

[food-pound onto wood [smash food))

used

Tai Forest

Cote divoire
<s

Wood-otlier Inut-hammer. wood hammer on earth

(stick

Budongo

Uganda

bodied

Wood-stone Inut-hammer, wood hammer

Lever open

criterion of

at start of rain)

Stone-wood {nut-hammer,

Pound

Kibale

witti petiolel

Stone-stone Inut-hammer. stone hammer on stone

Pound wood

meet the

apparent ecological

one other
Bossou

Wood-wood

w/ithout

o
o
o
o

o
o
o
o

One

variant

was absent

and another never


at the six

common

most studied

sites

listed tiere,

o
o

Nature, based on Wtiiten,

A.. e( at. 119991

NatureSI'):

682-685, used with

o
o

permission. For further


details, including less

studied sites (Mount

Assirili,

Lope], see V\/hiten, A., et ai.

1999.2003.

67

World Atlas

of Great Apes and their Conservation

DNA

markers and

has been noted

behavioral observations, found only one likely case

average wand

based on nuclear

extragroup

of

microsatelllte

among

paternity

may

incidence of 7 percent. Females

many males early in


later, when the likelihood

an

birtfis,

copulate

the receptive stage, but

with

of

conception

is

highest,

longer than

in

Tenkere, Sierra Leone, where the

was generally somewhat

length

Guinea, Senegal, and Tanzania, and

in

considerably longer than

in

however, the tool length

Cote dlvoire.'
is

longer tools used

attributes, with

In

Guinea,

determined by prey
higher-risk

in

they have been observed to copulate repeatedly only

contexts

with high-ranking adult males.'"

aggressive black Dorylus antsl. Here, two tech-

Communication

ants are eaten directly from the tool, and 'pull-

the ants' nest site or with the

at

(e.g.

niques are employed: direct mouthing', where the

Adult male chimpanzees often give loud pant-hoof

when

calls

they arrive at fruiting trees.

It

is

and the bundle

of

similar tools

tool

drawn through the hand

is

ants

is

then eaten.'" The use

has been noted

southwest

speculated that this behavior asserts the social

of

status of the caller, rather than being of benefit to

Cameroon,'"' '" indicating a wider distribution

the listeners,"'^' or that the purpose

the general use of tools than

and

to rally

is

maintain contact with the group while recruiting


allies

and associates.'" Younger males and females

The use

of tools to dip for driver

Uganda."

avoid attracting feeding competition and potentially

seen

join the

when

alpha male

in

they do, they

may

accommodate each

to

acquisition

acoustics, the backdrop of sounds


suite of wildlife,

and body size."

made
'^'

by the local

Other vocaliza-

tions include context-specific barks used

and snake alarms, and combinations

of

in

hunting

barks with

more

widespread among chimpanzee popu-

lations right across the geographical range of the

many

different

implements

purposes. Like humans, chimp-

used for a variety

of

anzees seem

be predominantly righthanded.'"

to

The tools thought

to

be most important

anzees are those used

in

chimp-

to

obtaining food and

inspecting their environment (including extracting,

proficient than

known

of

chimpanzees have

use,

TaV

some

minor context,

for

communities

different repertoires of tool

using far fewer tools than others.'^' The

chimpanzees have exhibited 28 out

of

42 tool-

use behaviors recorded throughout the range of the

compared with 17
Mahale (see Box 4.31.""

species,

The use by chimpanzees

in

Gombe and

13

in

ant-dipping

wands

playing,'^'

and there

Some games

are

while descending a slope, a behavior


in

the Mahale Mountains National

game

involves walking

backwards,

raking a pile of dead leaves along with both hands

and making a

lot of

use stepping

and

injury

noise.

Sierra Leone,

In

chimpanzees have been seen

sticks' (small sticks held

under the

'seat sticks' (sticks for sitting on), to avoid

from thorns encountered while feeding on the

flowers of kapok [Ceiba pentandra, Bombacaceae).'


In

Guinea, a chimpanzee

made from

cushion

was seen

sitting

[Carapa procera, Meliaceae), apparently


sitting

to

avoid

on wet ground.'"'

At Mount Assirik, chimpanzees used

sticks

on a

the leaves of the carapa tree

stalks to obtain termites


of

skill,

only from a particular locality, such as leaf-

Park.'" This

feet)

in

after acquiring the

not.'^'

pile pulling'

conspecifics or other species such as the leopard,

bodies.^' Different

Gombe

years for young

employed by her mother, whereas young

observed only

to

own

males

are cultural differences here too.

used for displays (including aggression against

cleaning their

five

develop the technique for termite-

Males spent more time

probing, and pounding]. Less important are tools

and communication! and,

took

be

termites about two years before males. Females are

males do

species, and involves

to

it

to

study of the

dipping.'" However, females had learnt to fish for

to that

Tool use
is

termite-dipping skills at

of

National Park found that

chimpanzees

learning.

in

and each young female uses a technique similar

acoustically distinct call types or drumming.^'

Tool use

mounds and

young chimpanzees spend time playing

All

pant-hoots vary somewhat geographically, which

such as habitat

was observed

Forest Reserve,

and learning from others, but there seem


sex-based differences

tentatively attributed to factors

ants

to perforate termite

other by giving acoustically similar calls.' The

is

in

previously known.

Equatorial Guinea, chimpanzees were

In

use sticks

to

in

then gather the termites with their hands.''

a chorus of pant-hoots and,

seem

was

for the first time in the Kalinzu

are generally quieter than adult males, possibly to

aggressive males." Younger males, however,

68

where the

through',

to

get

honey,

leaf

[Macrotermes subhyalinus],

and stones (probably as

Chimpanzee [Pan troglodytes]

break open the hard-shelled

hammers),

to

Adansonia

digitata (the

fruits of

baobab, Bombacaceael.'^

In

the CAR, a female chimpanzee used a large piece of


a

dead branch as a pounding

break

tool to

melipone beehive and obtain honey."

In

into a

the Gambia,

one chimpanzee was noted as sequentially using


a tool set comprising four different types of tools,

each with a different function,


a bees' nest

in

to extract

honey from

dead stump." Similar observations

made

have since been

Congo."

in

In

the Bwindi

Impenetrable Forest, Uganda, chimpanzees used


honey: a small stick for

of tools to obtain

two types

the tree cavities and subterranean holes used by a


stingless bee {Meiiponula bocandei]; and larger,

thicker tools to assist

foraging for honey of the

in

African honey bee [Apis

meWfera]

Chimpanzees also use

tools to crack nuts,

was documented

behavior that

in

Leone as

Sierra

long ago as the 16th century.^" At Bossou, they

use tools to open

oil

palm

nuts. Recent studies""'


in

Tool use for cracking

area could not be explained purely on the basis

nuts at Bossou, Guinea

there were

labovel and ant fishing

found that population-specific details of tool use


this

of

Malsuzawa Tetsuro

ecological differences - that

is,

cultural differences unrelated to ecology (Box

The techniques are learned when


three to

at

A. 31.

Ta'i

National Park,

Cote d'lvoire

the animals are

(left).

years old. the age at which juveniles

five

are also most likely to try unfamiliar foods

if

they are

offered them.

Chimpanzees

in

the Odzala-Koukoua National

Park used sedge (Cyperaceael stems

from the surface


to drink
in

of a

pond. The use

to

scoop algae

sponge

of a leaf

water from tree hollows has been observed

most long-term study

Bossou, chim-

sites.' At

panzees use folded leaves, most frequently from

Hybophrynium braunianum

((vtarantaceael, to obtain

drinking water from natural hollows


In

Tanzania,

some

in

trees."'

individuals have

been seen

using a nasal probe to induce sneezing, presumably


to clear out the

was once seen


in

nasal passage.

'

A chimpanzee

to insert a stick into a

narrow hole

hiding squirrel, which

a tree to rouse a

was

then captured and eaten."" Yet another Tanzanian

chimpanzee was seen wearing


a piece of skin

by a single overhand

whether

this

a necklace'

from red colobus;

was

it

knot but

tied by a

made

of

had been created


it

is

not

known
Ilka

chimpanzee.'^'

Herbinger/Wild Chimpanzee Foundalion IWCF)

lining.

Chimpanzees
trees.
to

build

They use

fairly

nests every night, usually

in

substantial branches or forks

form a framework and then bend and break side

branches

to

They may also

build different nests during

the day for resting; these are usually

Nest building

weave

a platform,

sometimes adding

sometimes on the ground." Up


be

built in a single tree

used varies

in

to

in

trees, but

10 nests

and the species

different regions. In

West

may

of tree

Africa, the

most commonly used species are the sassy tree

69

World Atlas

of Great Apes and their Conservation

Box U.k SEED DISPERSAL BY CHIMPANZEES

subsist on the poorer-quality end of this spectrum,

such as bark and herbaceous vegetation, but


The interaction between

plants and

fruiting

vertebrates that disperse their seeds


attracting

increasingly

is

chimpanzees are committed

it

is

conserve tropical forests

critical to retain

the frugivores that

The large mouth, robust

specialize

Chimpanzees process

Its

swallowing the seeds of

allocating

gape.

many

propelled

through

everywhere they have been studied. They ingest

chemical and

physical

much

larger

defecated whole and

than

smaller frugivores.

up

98 percent

to

absolute

amounts

terms.

time to

of their foraging

of fruit

and seeds per meal

both

Less than

population laround

percent

primate frugivores

in

Uganda

relative

in

percent

of all the

a large-bodied

the

of

represented by

is

for

an

esti-

seeds defecated."'

This high degree of frugivory

such

and

blomassi of

of the

chimpanzees, but they are responsible

mated 45.3 percent

fruit

mammal,

but

is

is

unusual

its

digestive system, which, like all the apes, features

a simple globular
for fermentative
is

stomach with no mechanism

digestion.

fruit

The digestive system

shared with the cercopithecme monkeys -

limits the

animal

to

eating foods that have relatively low concentrations


of toxins or digestion

and

fiber

some

In

inhibitors,

such as tannins

nature, such foods comprise fruits,

seeds, tender parts of plants, and animals.

The larger apes, the orangutans and

Seed dispersal
Reserve, Mali.

in

gorillas,

chimpanzee dung, Bafing

can

defined by
coarsely,

fruit

species

intact.

damage, and may be

large clumps."' For

in

Seeds

minimal

with

gut

some

chimpanzee's gut increases their germination

These factors

seed dispersers, which

Chimpanzees

amplified by their behavior

loaded

rate.

chimpanzees have

imply that

all

potential as

excellent

is

habitually

each day. infrequently dropping seed-

dung on the

forest

This foraging

floor

pattern facilitates long-distance seed transportation over a

wide area. This

crucial for tree species

is

such as Mimusops bagshawei (Sapotaceael. which


regenerates very poorly
of the

the Immediate vicinity

in

parent tree but produces viable seedlings

under other tree species.'"

Most chimpanzee-disseminated seeds are

macaques, guenons. mandrills, mangabeys, patas


monkeys, and baboons. This

the

need not

it

species, the passage of seeds through the

travel widely
in

consistent with

the lineage of the species and the design of

are

that

fruits of a particular size

in

show

fruit,

and manual

dentition,

chimpanzee mean

dexterity of a

disperse plants' seeds. Chimpanzees consistently


a year-round affinity for eating

and

frugivory

to

carnivory.

the attention of conservationists. This

reflects the thought that to


effectively,

the

either

upon by rodents or

preyed

succumb

pathogens (especially

to

insects,

fungi].

or

However.

these pressures are typically even greater for seeds

away from the area

that are not dispersed

may germinate

in situ,

dung

mostly

by

of the

consumed immediately

parent tree. The seeds not

or be dispersed secondarily
-

beetles

germinate

and

elsewhere. Germinating seeds face a cascade of


other destructive agents,

animals.

When

available,

however,

seed cohort does


lings

gap

the

some

finally establish to

As a by-product

of

canopy

forest

Is

fraction of the original

and subsequently grow

chimpanzees are

herbivorous

especially

in

become seed-

Into trees.

their foraging

effective

behavior,

seed dispersers over

long distances. This habitat-wide and year-round

broadcasting
species

IS

of

numerous seeds

of

multiple

prerequisite for the maintenance of

a heterogeneous forest. This

is

one reason

that

chimpanzees have been described as keystone


species

bers

in

may

forest ecosystems. Their decline In

therefore

num-

impair the composition and

structure of tropical forests.

James

70

V.

Wakibara

Chimpanzee [Pan troglodytes]

[Erythrophleum suaveoiens, Leguminosael and the


palm.'"

oil

Budongo Forest Reserve, the

the

In

chimpanzees

Cynometra

favored

particularly

a/exandn (Leguminosael. Here

it

was found

that day

nests were structurally simpler than night nests,

and were

same

Forest Reserve,

for feeding at the

used

built in the trees

height as feeding

where there are

the Kalinzu

In

activity.^'

relatively

many night nests are made


branches. In some places, nests are

carnivores,

few large
the low

in

also quite

frequently built on the ground, for example at


in

of

northern DRC, and

Bill

Nimba Mountains

the

in

Guinea."

Response to habitat disturbance


Chimpanzees are robust and adaptable animals
and have by far the widest geographical and
ecological distribution of any ape, perhaps of any

nonhuman primate in Africa apart from the


commoner species of baboon. One would therefore
chimpanzees

not expect
tive

be particularly sensi-

to

low or moderate degrees

to

such as might be caused by

disturbance,

logging

selective

ecosystem

of

light

by low

or patchy settlement

densities of farmers. Consistent with this, chimp-

anzee populations are known

logged forest, such as at Kalinzu

lightly

Reserve
is

survive well

to

in

Uganda." " Nevertheless, not

many

equally benign, and

significant

decline

all

studies have

logging

chimpanzee numbers

in

a
in

logged forest relative to comparable unlogged


areas,

for

example

in

Kalinzu,

'^^^"

Ituri,'"

and

Some groups

are

known

a time, in areas that have

almost

totally

travel

among

converted

to survive, at least for

been logged and then

to agriculture,

the few remaining

where they

small forest

patches and raid crops planted by local farmers."


Kibale National Park,

chimpanzees were found

nine out of 20 forest fragments

was their very


enabled them to use these
noted that

it

Chimpanzees
gorilla]

National Park

tend mutually to avoid contact^" [see Box

Baboons and chimpanzees


gether
In

one well studied example,

which are also eaten

baboons [Papio

the

local

(Leguminosael trees, which provide


in

and young leaves from September through

November.'"'
cultural

In

western Tanzania,

practices,

threats to the

were

survival

protected areas."

shifting agri-

uncontrolled bushfires, and

habitat fragmentation

of

identified

areas

habitat
at

higher

been observed

to eat

Red colobus

high

March, seeds

an unripe state by yellow

The baboons

moved

out of the park

and

as the major

populations outside

in

trees
floor,

altitude

alternative

food

they

had

at least

once

show

anti-

than

baboons.

in Tai

National Park

when chimps

are close, hiding

where they are shielded from the


and becoming

silent.

other

In

cumstances, however, they move closer


of

in-

range

their

chimpanzee group, which reacted by

exploiting

forest

fruits,

1975, reducing the ripe fruits available to

in

is

in

in

numbers and expanded

in

the Ugalla area, the survival of chimpanzees

July,

Mahale Mountains

cynocephalus]."'''

considerably after people

area

to-

those with a high calorie content,""

preferring

predation tactics

very important food: flowers

at

National Park, chimpanzees depend on ripe

In

threatened by the selective removal of Pterocarpus

8.11.

occur

often

formerly'" Mahale chimpanzees have

that

and they

the drier parts of the chimpanzee range.

in

fragments.

tinctorius

area of Guinea-Bissau.

differ

in

was

forest

Gabon], despite having similar

sources

it

1995, but

in

at Tombali,

typical of the coastal

keystone food resources

diets. Their

palm trees

example, the Lope

[for

In

home range

in

large

coexist with western gorillas {Goritta

some areas

in

creased

Budongo Forest Reserves.'"

Chimpanzee nests on

Interactions with other animals

in

Forest

shown

Claudia Sousa

to

cir-

groups

nearby Diana monkeys [Cercopithecus diana],

presumably because the


tinels for predators
floor"'^ In

latter are efficient

Uganda, red colobus have been seen

associate with several other species of

areas

same

sen-

approaching over the forest

of high

chimpanzee

monkeys

to
in

density, probably for the

reason." Chimpanzees also hunt guereza

71

World Atlas

of Great Apes and their Conservation

Table i.2

A summary

chimpanzee

of population data for the

Western

Central

Eastern

Nigeria-

Cameroon
200-500

Angola
Benin

extinct

extincf

Burkina Faso

200-500

Burundi

Cameroon

000-39 000

31

CAR

3 000-5 000

unknown'

800-1 000

Congo

10 000
8 000-12 000

Cote d'lvoire

DRC

70 000-110 000

extincf

Equatorial Guinea

000-2 000

27 000-6^ 000

Gabon

Ghana

300-500

Guinea

8 100-29 000

600-1 000

Guinea-Bissau
Liberia

000-5 000

Mali

600-5 200
2 000-3 000

unconfirmed^

Nigeria

Rwanda

500

200-AOO

Senegal
Sierra

Leone

500-2 500

200-400

Sudan
Togo

extinct

Uganda

4 000-5 700

United Rep. of Tanzania

70 000-117 000

Total

a 'Unknown' indicates that

As the

table shows, the

Reserves

in

it

not clear

is

how

21

large the population

Nigeria-Cameroon chimpanzee

Cameroon have

is

is;

500-2 500

76 000-120 000

000-56 000

'unconfirmed' indicates that there

by far the most rare. The Ejagham and

may

000-8 000

not be a population.

Takamanda

Forest

also been referred to as having a 'significant population' of the Nigeria-Cameroon

chimpanzee.'"
Data compiled by Butynski,

TM

(20031:^^

see Chapter 16 for later estimates for Cameroon,

Ghana, Mali, and Nigeria as well as further details

ICoiobus guereza]

in

latter is significantly

Uganda, and the density


lower

in

chimpanzee

of the

activity

centers than outside them.'"' Guerezas appear


react to the presence of

way from
their

chimpanzees

in a

to

make

escape along the ground.'"

chimpanzees

latter

in

found nesting inside a night

nest of the former"^


In Tai

National Park, 29 interactions between

leopards and chimpanzees were observed from

985

chimpanzees were

observed

killed,

with the leopards apparently being the main

capturing and toying with western tree hyraxes

cause

of mortality in the area.'"

[Dendrohyrax dorsalis], but did not eat them and

found

in

appeared not

probably been killed by a leopard." However, the

In

Guinea,

Similarly, a

as a

toy,

to regard the

hyrax as a prey animal.'"'

Mahale chimpanzee treated a squirrel

making play faces during the encounter

after the animal was dead.'"' A


more benign association between chimpanzees and

and giving up shortly

72

Uganda, with the

to 1990. In these, at least four

were

populations

Thomas's galago [Galago thomasi] was noted

different

red colobus; they remain quiet and

of national

A dead chimpanzee

the Petit Loango Reserve,

attacks do not all run one way.

In

Gabon had also

Tanzania, a group of

about 33 chimpanzees surrounded a mother leopard

and her cub


the

in

their den,

cub,'" while

in

and dragged out and

Uganda

killed

small group

of

Chimpanzee {Pan troglodytes]

chimpanzees chased

oft a leopard.^'' In

Mountains National Park, evidence

was found

chinnpanzees

the Mahale

of lions eating

1989.""

in

Unless habituated, chinnpanzees usually react


with fear towards hunnan beings.
recently

Gombe

in

National Park (20001,

Kibale

attacked and killed

On occasion,

human

chinnpanzees have

Most primatolo-

infants.

chimpanzees are driven by

gists believe that the

same

infan-

to attacks

on the

predatory instincts rather than by the


ticidal

urges that sometimes lead

Whatever the motivation,

offspring of rival males.

these rare events are devastating

damaging

affected and

nnost

National Park (2002) and near

the families

to

conservation efforts.

to local

POPULATION
Status
It

is

number

estimate the current

to

difficult

chimpanzees because recent information


from many areas and nothing

some. Data are summarized

at alt

is

of

lacking

known about

is

Table A.2, and indi-

in

cate an estimated total population size of between

172 000 and 301 000 chimpanzees

2003."

in

Trends

As with other

forest animals,

populations. Attempts have been


overall

in

Western chimpanzees
climbing, Cote d'lvoire.

made

to

Lk;

to

estimate

numbers by applying population


known

values at
suitable

assess

chimpanzee

is difficult

it

population size and monitor trends

habitat

Herbinger/WiLd Chimpanzee Foundation IWCF)

Elizabeth A. Williamson

the remaining area of

sites to
in

density

the species'

range. The total

number of chimpanzees in 1987 was estimated


at 151000-235 000."' and in 1989 at U5 000228 000.'" The figures

in

Table 4.2 for 2003 suggest

may

that these previous totals

estimates.

It

have been under-

has been argued that there were some

2 million chimpanzees around 1900, and


1

is

million as recently as 1960."

consistent with

much

else that

during the 20th century,

more than

decline on this scale

happened

in

Africa

including widespread

deforestation, the expansion of farming and infra-

structure at alt scales, increased access to military


firearms, and

Trends
difficult to

human
in

population growth.

individual

assess because

countries are similarly

many

previous estimates

were probably underestimates. However, an example


of decline

quoted by Butynski'"

claimed that

from 20 000
1987.'"

In

in
in

Sierra

is

notable:

it

was

Leone the population dropped

the late 19th century to 2 000

Gabon, the combined population

chimpanzees and

gorillas declined by

in

of

more than 50

percent between 1983 and 2000 due to the increase

This infant central

chimpanzee
in

pack

is

strapped

the traditional back


of the

poacher who

killed its

mother,

Cameroon.

73

World Atlas

of Great Apes and their Conservation

ORPHAN

Box 4.5 REINTRODUCTION OF

CHIMPANZEES
1989, Allette

In

Jamart set up the project Habitat

Ecologique et Liberte des Primates (HELP Congo],


the

aim

of

which was

reintroduce chimpanzees

to

rescued from the bushmeat trade


environment. These

to their natural

are

activities

based

the

in

Conkouati-Douli National Park, Congo.

From the
the
to

the project aimed to follow

start,

lUCN Primate Reintroduction Guidelines, and

evaluate the reasons for successes and/or

lures

the short,

in

stages

of

medium and

HELP

the

project are

long term.

fai-

The main

summarized

below.
ns/HELP

three islands were identified

1989:

the

in

Conkouati lagoon as suitable for the establishment

of

the

HELP orphan

1990: the sanctuary received

sanctuary;
its first

group

group

first

chimpanzees was

of

the fitting of radiotransmitters;

in

of

chimpanzees from

1996-2000: a

made

by Marc Ancrenaz

individual

collaboration with the Centre International

total of

seven releases

chimpanzees

37

of

males and 27

(10

females! have taken place Isee tablel.

Recherches Medicates de Franceville

ICIRMFI, Gabon;

HELP

199A: an evaluation by Caroline Tutin on be-

chimpanzees, as they are well received by wild

half of

lUCN was

carried out to establish the

release possibilities for the chimpanzees

in

the Conkouati area Itwo sites were identified!;

1995

second medical examination was

earned out on the group, again by Marc

Ancrenaz

showed

of

CIRMF, the results

that they

were

which

of

free of disease

and

ready for release;


1996: a second
Tutin

out

evaluation, by Caroline

and the botanist Paul

in

males and can go on


the

wild.

Sita,

was

carried

the potential release sites, resulting

in

reproduce successfully

to

The capture

of

chimpanzees

sanctuary, their transport to the release


their introduction into a

can be sources

of intense stress for the

the

journey

to

the release site and

at Triangle Island.

when

awake

The cage door

chimpanzees

that

Is

have

99 percent decline

In

MInkebe Forest,

the northeast, since 1994,

in

chimpanzees was recorded

was an Ebola epidemic."^ A sudden


was

also noted

since

the

in

Gombe

mid-1990s,

In

in

when

population

National Park,

owing

combination of hunting and disease.'^ "

was noted

Gabon, a

In

to

Sudan,

1964 that the species "appeared

thrive particularly well in

were reported
individuals,

to

SW Sudan

move around

sometimes

cage

their

opened only
been

In

from where they

bands

30

of

more,"''" but by 1988

to
It

40

was

stated that the "species could be considered highly

endangered

If

Records

not already extinct


at

picture, ranging

it

Kllum-ljim forest

to

in

already

are fully conscious.

in

animals. To

be close to the cage and the

new chimpanzees

the country.'" Also

and

habitat

animals are unconscious through the

by rivers adjoining Conkouati National Park;

In

site,

agents ketamine and medetomidine are given, so

to

logging and hunting, and the spread of Ebola

the

of the anesthetic

released are seen

hemorrhagic fever

in

at

new and unknown

reduce this stress, a combination

that

lUCN

prefers to concentrate on releasing female

selection of the 'Triangle', an area surrounded

Tanzania

74

1996: the

of

1992: preliminary medical checks of captive

de

decline

year old male central chimpanzee leaving the

released, after a final medical evaluation and

the sanctuary were

there

six

cage during his release.

chimpanzees from Pointe Noire;

and semicaptive groups

in

International

individual

from
in

in

sites

the country."'"

show

varied

local extinctions (e.g. at the

northwest Cameroon

19881'" to stability, or even recovery.

In

in

1987 or

the Monte

Chimpanzee [Pan troglodytes]

ranging over quite a wide

Post-release monitoring
Following

their

release,

the chimpanzees are

was

monitored

daily

from nest

to nest for a period of

ti'lay

weeks

acclimatization that varies from several

months, depending on the


death rate

is

individual.

14 percent, the survival rate

percent, and the disappearance rate


If

they have been attacked

to

To date, the

is

is

by wild males, the

released males are then followed from morning

through

to evening.

daily basis, but

when

in

The females are located on

tend to leave the monitoring area

'estrus'.

It

thought that they socialize

is

Massabi was reobserved

for

months

six

returned with her baby; Rosette

months but then spent


chimpanzees; Matalila
returned

to stay

then

but

was absent

for 18

5 days with the monitored

left

for

months and then

with her childhood group; Massabi

and Mossendjo were found two and a


after release in a

marshy

area,

half

Only Massabi
it

was

lost in

in

January 2004,

in

the

male.

of a wild

follow-up study of the released chimpan-

zees has highlighted that male chimpanzees


cannot be released where wild chimpanzees
as they are

be

to

likely

killed

by the

exist,

existing

Chimpanzee release was smoother

population.

when animals were

anesthetized tor transport,

and released soon after recovering consciousness.


While reintroduction

with wild males during these periods. Bonnie, for

example, was absent

when

no longer be found.

2003, the pair could

company

62

24 percent.

territory.

with a radio collar;

fitted

is

not the only possible

solution to the overpopulation of

sanctuaries,
tool.

it

has proved

of

in

priority for

chimp-

protect their habitat and

is to

reduce the pressures

chimpanzees

be a potentially useful

mam

Nevertheless, the

anzee conservation

to

hunting.

months

where they were

Jamart and Benoit Goossens

Aiiette

Release methods and problems encountered

Date

Number
(male,

1996

1^;

released
female,

Transport to point

Situation prior

Problems following

of release

to release

release

Fl

floating cage;

5I1M, 4FI

floating

none

cage

animals conscious
1997

floating cage;

2I2FI

direct release

animals panicked

release cage

animals stressed

and escaped

animals anesthetized
1997

boat;

8 I2M, 6FI

and panicked by

animals anesthetized

captive conditions

1999

animals stressed

release cage

boat;

5 I2M, 3FI

and panicked by

animals anesthetized

captive conditions

2000

4|1M,3FI

floating

boat;

animals anesthetized

until

none

cage

conscious at

release site

2001

into

2001

12(3M,9F|

boat

an existing group

floating

awake

1994, for

Alen National Park, Equatorial Guinea,

in

example, chimpanzees were found

common

to

be

none

released directly

boat

IMl

all

at

cage

none

until

launch

site

Threats
Estimates

of extinction risk for

chimpanzees are

over the park and were not apparently threatened

largely based on the observed loss or modification

by hunting as they had been

of

in

previous years."

In

Kibale National Park, Uganda, two sets of censuses

were carried out


1975-1976 and

in

old-growth forests during

1997-1998;

it

was found

chimpanzee populations had declined only


nificantly over this period.'"

their

also,

in

habitats,

the

case

on rates
of

of

exploitation,

and

geographically restricted

populations, on the risks that are inherent to a

that

small range

insig-

Commission

size.

In

2000, the Species Survival

lUCN-The World Conservation


Union categorized the chimpanzee species as
of

75

World Atlas

of Great Apes and their Conservation

Box U.b CHIMPANZEE HABITUATION FOR


TOURISM

in

chimpanzees,

Habituating

view.

however,

is

quick or easy task, due to their ranging

not a

behavior and their fission-fusion society The size of

National Park

Kibate

gazetted

1993 and

in

managed

is

Wildlife Authority lUWAl.


of

western Uganda was

in

by the

comprises about 795 km^

It

moist evergreen forest, colonizing forest, papyrus

swamp, and

exotic softwood

plantation, and

surrounded by a dense human population.

most important
in

Uganda

over a quarter

400 individuals, homes

the country's population.^" Three

and Ngogo,

for behavioral

Kanyanchu,

for tourism.

The potential

for

1980s.

to

primate tourism within

home range

be located and the presence

offering forest

the late

1991.

walks with the chance

of viewing

chimpanzees varied considerably,

however, a project
level of

was

1997

initiated in

habituation and gain a

more

to raise the

consistent

viewing rate for tourists. These were the aims of the


Kibale Primate Habituation Project, a joint venture
of

UWA and

of individuals within a

chimpanzee

inevitably intermittent.

Kibale Primate Habituation Project

From 1997

the project worked alongside 12

to 2001,

Ugandan rangers to maintain dawn-to-dusk contact

A team

with the chimpanzees.

two rangers

of

chimpanzees early

by returning to their nest

site

in

typi-

the morning

or popular feeding

trees, or by being guided by their calls. Habituators

stayed with the chimpanzees, collecting baseline

ecology, and range use. Observer protocols designed

In

to tourists,

of finding

is

of fruit in

chimpanzees and other primates. As the success


rate

number

large

community

data on party composition and interactions, feeding

opened

Visitors Centre

individuals

chimp-

to allow

their preferred food trees to be monitored.

Kanyanchu

same

habituation and tourist visits the

are not consistently followed, and contact with a

cally located the

system was established within the

trail

chimpanzees' core

anzees

research purposes; and

was recognized during

Kibale National Park

the

is

the

communities are habituated: Kanyawara

the

of

of

is

parties can vary from two to over 50

individuals of different age-sex classes. During

chimpanzees

habitat for eastern

Uganda and, with over

It

chimpanzee

the Jane Goodall Institute-Uganda.

tourism were adopted and habituators

for gorilla

took care not to


staring,

show threatening behavior such as

sudden movement, or close proximity The

well maintained

access deeper
trained

in

trail

system was expanded

into the

home

to

allow

range. Rangers were

chimpanzee behavior, ecology, data


and guiding, and were well equipped

collection,

with binoculars, compasses, backpacks, uniforms,


boots, and

ram gear

enable habituators

All

to

rangers carried radios

communicate the

to

location of

the chimpanzees to those guiding visitors.

Habituation for tourism

The

project

aimed

to

achieve a level of habituation

balanced the need for chimpanzees

that

relaxed

enough

in

the

presence

of

behave naturally while maintaining a

to

humans

to

sufficient de-

the

four years,

majority

chimpanzee

of

individuals reached a level of habituation that

permit them

to

be followed

all day.

would

often at close

gree of wariness to prevent aggressive encounters.

quarters. The monthly average success rate for


visitor

level of habituation

is

the tourist experience,

typically

of critical

importance

as wild chimpanzees

disappear quickly at the sight of humans.

Observations

of

some

of

the

most

interesting

aspects of chimpanzee behavior, such as the use


tools,

is

Endangered,
the wild

of

correlated with the degree of habituation

and the length

in

i.e.

of

time that the animals can be kept

facing a very high risk of extinction

chimpanzee populations. Light

groups viewing chimpanzees rose from 61

percent

in

individual

1997

to

88 percent

chimpanzees were

Kanyanchu

is

one

ties in the wild, with

also

selective

logging

causes only temporary disturbance and may not

in

2001. Over 60

identified

of the largest

and named.

known communi-

22 adult males. The project

compiled a database with

information

on

demography, range use. feeding patterns, social

greatly reduce the forest's carrying

chimpanzees unless

the near future.

Threats multiply one another's impacts on

76

Project achievements

After

The
to

in

be

particularly

capacity for

favored

tree

species are lost from the biota. Increasingly intense


logging and repeated re-entry logging, however, will

cause mounting disruption

to the forest

ecosystem

Chimpanzee [Pan troglodytes]

and associations. The long-term

hierarchy,

chimpanzees and provide

individual

interpretations

community, and

chimpanzee track-

skilled rangers,

Kanyanchu

ing in

The project also helped

diversify

in

following

chimpanzees

and witnessing the chimpanzees'

Uganda

for viewing wild

the subsequent increase


region,

full

range

day

all

of daily

Kanyanchu has become the most popular

activities.

site in

tourism

the unique opportunity to join the

team

IS

important

disease transmission

chimpanzees. With

numbers

effects

and

direction,

or population viability

duration

behavior are

The

positive

UWA

local people.

tourism on chimpanzee behavior,

of

understood, so the
day,

to avoid

maximize

to

and minimize the negative impacts on the

ecology,

of
all

is

currently

optimum number
visits,

little

of tourists

per

observer

or appropriate

a matter of educated guesswork.

and Makerere University are currently

collaborating to monitor the impacts of tourism on


the
will

enterprises outside the park boundaries, such as

each
is

contact

the apes, to reduce

to

chimpanzees, the environment, and the

community-run tourism and conservation

tourist

in

aggression. The goal

to the

in

manage human-chimpanzee

to

so as to minimize stress

The impact

a rewarding experience.

is

activities, offering

habituation

their

to

forest, a large habituated

With a stunning

visitors.

interesting

chimpanzee behavior

of

local

named

can identify the majority of the

staff

field

Kanyanchu chimpanzee community. The


be used

to revise

results

observer protocols,

courage their adherence, and

to

to

en-

design long-term

monitoring systems aimed at detecting changes

the Bigodi Wetland Sanctuary, have flourished.

in

the behavior, health, and environment of the chimp-

anzees before any

Conservation and challenges for the future

irretrievable

The benefits accruing from chimpanzee tourism

communities, and reducing

local

and

poverty, poaching,

such important

forest

of

levels

encroachment. Despite
remain

benefits, challenges

and

is

done.

Ajarova

Lilly

management

include revenue generation for park

and

damage

Julia Lloyd

A tree house used

for habituation in Kibale

National Park, Uganda.


Julia Lloyd

for the

future, notably the successful coordination of differ-

ent tourism activities, adherence to observer protocols,

and monitoring

of habituated

substantial level of chimpanzee

chimpanzees lose

however, they could


visitors,

and

habituation

followed

sense

all

chimpanzees.

experience

tourism

high-quality

requires

habituation;

if

of fear of

pose a threat

to

humans,
rangers,

Once the optimum

local people.

level of

reached, the chimpanzees must be

is

daily

allow them to

to

be located and

monitored, which requires numerous well trained


staff

and

effective on-site

The use

of

due

sensitive issue

to their

original regulations for

based on those

pitfalls.'^'

and

its

not yet known.

its

shine,

opening

is

humans. The

chimpanzee tourism were

own

set

of

chimpanzee

challenges and

long-term impact on chimpanzees

As with gorilla-focused tourism,

which chimpanzees are a

integrity,

tourism

for gorilla tourism, but

tourism offers

of

in

Endangered status and

evolutionary relationship to

close

is

management.

chimpanzees

it

and increasing

up
its

part,

to drying

degrading

it

its

winds and sun-

vulnerability to forest fires.

Increased access to the logged-over area

along

logging

roads will

encourage entry by

hunters, especially
trade

in

bushmeat

where there

las there

is in

is

a commercial

much

of

West and

Central Africa), so the effects of hunting almost


inevitably
oil

compound those

of logging. Ivlining

and

extraction can have simitar effects on access, as

77

World Atlas

The boundary

of Great Apes and their Conservation

of Kibira

National Park, Kabarore,

Burundi. Here, as

elsewhere, chimpanzee
habitat

is

threatened

by encroaching
agriculture.
Geoffroy Citegetse

natural ecosystems.

well as locally destroying

Increased access also leaves the forest open to

the region, and the assumption that

have been

Bushmeat

invasion by settlers, leading to further hunting and

also to the fragmentation

of

the

by an

forest

in

each

of

often a

is

chimpanzees

these orphans.

major source

of dietary

the meat-eating cultures of West and

expanding and eventually coalescing mosaic

of

Central Africa, and sometimes also has perceived

Chimpanzees are bound

to

magical or medicinal benefits. Although hunting

farms and
find

it

villages.

increasingly difficult to survive

in

such a

more so as the reduced and


fragmented populations come into more frequent
contact with humans and become increasingly
landscape,

vulnerable

the

human

to

chimpanzees,

diseases. Hence the survival of

many

that of

lil<e

other species,

is

may occur

at sustainable levels locally,

with logging and mining because food

human communities

often

favor bushmeat.

people from other regions. The impact

now widespread,

human use

parallel with greater access to

moist forests.

With deforestation so far advanced

remnant

persist.

The fragmented populations

and western subspecies


marily located

and national

in

of

remnant

parl<s;

in

West

tracts of primary rain forest


of the

eastern

chimpanzee are

forest,

game

pri-

reserves,

unauthorized hunting, logging,

mining, and farming are

common

in

many nomi-

nally protected areas.

Hunting

of adults for

on populations and

is

threat to the species.

bushmeat has an impact

an important and increasing

report released

in

2004 has

is

markets are being developed

widely sold
in

infant

in

adults. In

some

Kouilou Basin

it

conclusion

with extinction within 17-23 years. This

was based on

orphaned chimpanzees arriving

at

to serve rising

in

New

demand

areas, however, for example the


is

not hunt-

ed for meat and

is

Not

intentional, however; trapping

all

hunting

is

consequently less threatened."

target prey species.


11 of

can

chimpanzees even where they are not the

injure

Limb deformities were found

52 chimpanzees
all of

living in

them
in

The dual impacts

number

of

bushmeat hunting

sanctuaries

in

gorilla

the increasing

remote areas.

Congo, the chimpanzee

in

Reserve, nearly

threaten

bushmeat

often associated with hunting of

is

and leg-hold traps set

to

of

increasing rapidly

and regional markets, and trade

local

chimps

panzee, bushmeat hunting alone

sufficient

Civil

from urban populations, chimpanzee products are

estimated that for the Nigeria-Cameroon chimis

increases

required to

also tends to increase hunting, often by

conflict

hunting

of tropical

it

is

maintain large labor forces, and because colonizing

threatened by the whole process of advancing

Africa, only

78

protein

for

l<illed

in

the

in

Budongo Forest

attributed to wire snares

the forest.
of

the

Ebola virus and

the heartlands of the western

and chimpanzee range

in

the

Congo Basin

Chimpanzee {Pan troglodytes]

and may have already greatly

unquantified

are

of both species.'"

reduced populations

new Ebola

of a

killed

strain

in

the Tai Forest, Cote d'lvoire

2 chimpanzees, about a quarter of the group

under

study."' ""

park's

chimpanzees died

most

the

An outbreak

is

It

not

known how many


in

Ebola

total.

of the

by far

is

disease affecting the African

virulent

great apes, but others have also had significant

and

Wallis

impacts.

occurrence

of

disease

National Park, Tanzania,


'polio-like' virus,

summarized

Lee^"
in

chimpanzees
noting

the

Gombe

in

incidences of a

pneumonia and other respiratory

diseases, and scabies. They discussed the posthat these diseases

sibility

had been contracted

from humans and suggested various improvements


in

the deaths of

all

targeted

party

other chimpanzees present

hunters. Although

by the

concern has been expressed

in

the

in

much

the past over such

uses and the possible impact on wild populations,

most areas

this is

habitat loss

amount

in

thought to be a lesser threat than

and the bushmeat trade. A reasonable

of conservation attention

rescue and rehabilitation

of the

both for their intrinsic value and

an educational opportunity out


disaster Release

schemes

for

focused on the

is

orphans themselves,
order

in

to

create

of a conservation

rescued chimpanzees

are not as far advanced as they are for orangutans,


but are

more advanced than

successful

trial is

ongoing

in

National Park, Congo (see Box

for

One

gorillas.

the Conkouati-Douli
4.51.

health standards to help combat the problem.

Also

in

chimpanzees

Tanzania,

are

affected

CONSERVATION

seasonally by intestinal nematodes, particularly

Chimpanzees are the most abundant and wide-

Oesophagostomum stephanostomum. which can

spread

result

in

secondary bacterial

infection,

diarrhea,

of the great apes, with a total wild population

estimated

to

be up

to

300 000 individuals. Most

live

severe abdominal pain, and weakness, sometimes

outside protected areas, where they are vulnerable

leading to death. Disease transmission and Ebola

to

are discussed further

in

Chapter

Finally, the live-animal

of

for

infants

industry,

the

pet

trade involves capture

trade,

the

entertainment

and international biomedical business.

Like hunting, this

capture of

is

illegal in all

disturbance of their forest habitat by logging; to

habitat destruction by settlement,

13.

range states. The

an infant chimpanzee typically involves

and

to

fire,

and farming;

hunting that supplies the increasingly en-

trenched and powerful bushmeat trade. Meanwhile,


their

fragmented populations are becoming increas-

ingly subject to disease

more

outbreaks as they

come

often into contact with people. The detail of

Table 4.3 Conservation success scores for protected areas

Protected area

Chimpanzee

Conservation

subspecies

success score'

Cameroon

Korup National Park

central

3.0

Cameroon

Dja Faunal Reserve

central

3.0

CAR

Dzanga-Sangha

eastern

3.5

Congo

Odzala National Park

central

5.0

Cote d'lvoire

Marahoue National Park

western

2.0

Cote

Tai National

western

3,8

d'lvoire

Park

DRC

Itun Forest Reserve

eastern

3.5

Equatorial Guinea

Monte Alen National Park

central

5.0

Gabon

Lope National Park

central

3.0

Ghana

Bia National Park

western

2,0

Nigeria

Cross River National Park

Nigeria-Cameroon

3.0

Uganda

Kibale National Park

eastern

3.9

United Rep. of Tanzania

Mahale Mountains National Park

eastern

4.0

a Consen/ation success scores derive from qualitative questionnaire assessments

in wtiicii

indicates failure

and

5 indicates very successful.

Data compiled by Strutisal<er. T T. e( al 120051

79

World Atlas

of Great Apes and their Conservation

conservation efforts for chimpanzees varies over

broad range, as outlined

tfieir

country profiles

As these

in

Chapter 16

profiles

the

in

this

3.5;

their protected areas

of all four

In

protected areas

in

median was

suggests

that,

overall,

informed observers were reasonably confident that

volume.

of this

show, populations

chimpanzee subspecies occur

the

relevant

were working.

decreasing order of significance, the main

factors contributing to an increase

across their range. These protected areas occupy

success scores were considered

in

conservation

to be:

tens of thousands of square l<ilometers of forest


that range-state

governments have chosen

to set

1.

a positive public attitude;

aside for wildlife protection. The central issues that

2.

effective

emerge

which these

3.

large protected area size;

areas are managed, the challenges that they must

4.

low

overcome, and ultimately the security

5.

the presence of nongovernmental

are

investment

scheme
to

is

the

in

effectiveness with

of the public

conservation that they represent.

being developed

from tourism (Box

sample

in

the vicinity;

ecological continuity

On average, the lowest scores were obtained

of 13 protected

is

one

of

West African protected areas,

an Africa-wide

areas containing chimpan-

zees that have been analyzed from the point

conservation effectiveness."^ These are listed

and the presence

the

overall

approach was used

programs

their area of distribution,

dismal

may

overall, intractable

public

that

investment compromised

and debt, the outlook

mean

uncertain at best.

13 protected areas

all

was

3.4,

grounds

be

and

their

prospects.
for

patchy

human populations rising


poverty in many locations, and

would have allowed a quantitative evaluation. The


score for

conservation

optimism, although with

the absence of

in

protected area-wide monitoring

in

more

Elsewhere, there

to 5, with

being a failure and 5 being very successful. This

qualitative

is

two West African subspecies, the greater

deforestation

with the area concerned. Respondents were asked

score the protected area on a scale of

dense human populations

of

generally consistent with the lower abundance of

and protected area managers familiar

of scientists

for

their

possessing a bushmeat-eating culture. This

in

which represents the qualitative opinion

for each,

reflecting

greater ecological isolation, greater accessibility,

view

of

Table 4.3, along with a 'conservation success score'

population density

organizations; and
6.

A.6].

Kibale National Parl<

to

human

supplement public investment with funds accru-

ing

of

Kibale National Park

in

law enforcement;

and

for

by corruption

chimpanzees remains

FURTHER READING
Chapman,

C.A..

Onderdonk, D.A. 11998) Forests without primates: primate/plant codependency. American Journal of

Primatology i5lM 127-U1.

Dominy,

N.J.,

Human
Goodall,

J.

Svenning, J-C,

Li,

W-H.

120031 Historical contingency

in

the evolution of primate colour vision. Journal of

Evolution 44: 25-45.

11986)

Ttie

Cfiimpanzees of Gombe: Patterns of Bef)avior. Harvard University Press, Cambridge,

Massachusetts.
Goodall,

Grubb,

J.

P.,

11990) Ttirougt) a Window:

Butynski, TM., Gates,

My Ttiirty Years

J.F.,

with the

Chimpanzees of Gombe. Houghton

Bearder, S.K., Disotell, TR., Groves,

C.P.,

Mifflin

Company, Boston.

Struhsaker, T.T 120031 Assessment of

the diversity of African primates. International Journal of Primatology2U[b\: 1301-1357.

Kormos,

R.,

Boesch, C. 12003) Regional Action Plan for the Conservation of Chimpanzees

in

West Africa. Conservation

International, Washington, DC.

Kormos,

R.,

Boesch, C, Bakarr,

M.I.,

Butynski, TM., eds 12003) West African Chimpanzees: Status Survey

and

Conservation Action Plan. lUCN/SSC Primate Specialist Group. lUCN, Gland, Switzerland.
Kortlandt, A. 11983) Marginal habitats of chimpanzees. Journal of Human Evolution 12

Lonsdort,

E.V.,

Eberly, I.E., Pusey, A.E. 12004) Sex differences

McGrew, W.C, Marchant,

L.F.,

Nishida, T, eds 11996)

in

learning

in

131:

231-278.

chimpanzees. NatureiZS: 715-716.

Great Ape Societies. Cambridge University Press,

Cambridge, UK.
Mltanl, J.C, Watts, D. 11999)

Demographic influences on the hunting behavior

Physical Anthropology 1 09: 439-454.

80

of

chimpanzees. American Journal of

Chimpanzee [Pan troglodytes]

Parr, L.A., de Waal, F.B.M. (19991 Visual kin recognition

Reynolds,

Reynolds,

V.,

in

chimpanzees. Nature399: 6i7-648.

Budongo

119651 Chinnpanzees of the

F.

Studies of Monkeys and Apes. Holt, Rinehart and Winston,

Forest.

New

In:

DeVore,

I.,

Chimpanzee and Red Colobus: The Ecology of Predator and

Stanford, C.B. 119981

Primate Behaviour. Field

ed..

York. pp. 368-^24.

Prey. Harvard University Press,

Cambridge, Massachusetts.
Stanford, C.B. ,Wallis,
in

Gombe

Struhsaker,

J.,

Matama,

Goodall,

H.,

J.

11994) Patterns of predalion by

T.T.,

Struhsaker,

P.J.,

Siex, K.S. 12005) Conserving Africa's

possible solutions. Biological Conservation 123

Whiten,

Goodall,

A.,

McGrew/, W.C, Nishida.

J..

Horner,

A.,

V.,

111:

forests:

problems

9ii:

in

213-228.

protected areas and

45-54.

Reynolds.

T.,

ram

V.,

Sugiyama,

Y.,

Tutin, C.E.G.,

Wrangham,

R.W., Boesch,

chimpanzees, Wa(ure399: 682-685.

C. (1999) Cultures in

Whiten,

chimpanzees on red colobus monkeys

National Park, Tanzania, 1982-1991. American Journal of Physical Anthropology

C, Marshall-Pescini,

Litchfield,

S.R.J. (20031 Cultural

panthropology Evolutionary Anthropology

12121:92-105.

Wrangham,

Chapman,

R.W.,

Tropical Ecology

Wrangham,

Chapman,

C.A.,

(1994) Seed dispersal by forest

L.J.

chimpanzees

in

Uganda. Journal of

W: 355-368.

R.W., McGrew/,

W.C, de Waal,

RB.I>^.,

Heltne, P.G., eds (1994)

Chimpanzee Cultures. Harvard

University

Press, Cambridge, Massachusetts.

MAP DATA SOURCES


Map

4.1

Chimpanzee data are based on the

Butynski, T.M. (2001) Africa's great apes.

Stevens, E.R, Arluke,

A., eds.

following sources, with updates as

In:

Beck, B.B., Stoinski,

T.S.,

in

the country profiles

Hutchins, M., Maple,

T.L.,

in

Norton,

Chapter
B.,

16:

Rowan,

A.,

Great Apes and Humans: The Ethics of Coexistence. Smithsonian Institution Press,

Washington, DC. pp. 3-56.


Butynski, T.M. 12003) The chimpanzee

Kormos,

R.,

Pan

Boesch, C, Bakarr,

troglodytes, taxonomy, distribution, abundance,

M.I.,

Butynski, TM., eds.

and conservation

status.

In:

West African Chimpanzees: Status Survey and

Conservation Action Plan. lUCN/SSC Primate Specialist Group. lUCN, Gland. Switzerland, pp. 5-12.
For protected area and other data, see Using the maps'.

ACKNOWLEDGMENTS
Many thanks

to Colin

Groves lAustralian National University), Phyllis Lee (Cambridge University), John

College, City University of

comments on

New

York!,

and David Woodruff (University

of California,

San Diego)

R Dates

the draft of this chapter

HELP Congo,

discussed

in

Box

4.5.

works

in

collaboration with the authorities of

Congo

at the Ministry of Forestry

and the Environment and with the Wildlife Conservation Society [WCS-Congol. HELP Congo wishes
sponsors:

US

Columbus Zoo and Aquarium;

Fish and Wildlife Service; Cleveland Zoological Society;

International Primate Protection League; Arcus Foundation;

Bardot; Fondation Bourdon; Societe Protectrice des Animaux;


Gorilla; Air

(Hunter

for their valuable

to

thank

Pan African Sanctuaries Alliance; Fondation

One

Voice; Beauval Zoo; Amneville Zoo; La

Gabon; and Cardiff University; as well as the Congolese assistants who

daily follow the

its

Lincoln Park Zoo;


Brigitte

Barben Zoo;

chimpanzees

in

the

forest of the Conkouati-Douli National Park.

The chimpanzee habituation


Grants

Management

Unit, the

project

in

the Kibale National Park, discussed

in

Box

4.6,

was funded by

the

USAID

Jane Goodall Institute-Uganda, Cleveland Zoo, North Carolina Zoo, Barclays Bank

(Kampala!, British Airways, Discovery

Initiatives,

and TOTAL lUG)

Ltd.

AUTHORS
UNEP World

Tim

Inskipp,

Box

4.1

Box

4.2 Alison Surridge,

Box

4.3

Andrew Whiten,

Box 4.4 James


Box 4.5

Conservation Monitoring Centre

Craig Stanford, Jane Goodall Research Center, University of Southern California

Aliette

School

of Biological Sciences, University of East Anglia

Scottish Primate Research Group, University of St

Andrews

Wakibara, Tanzania National Parks/Kyoto University

Jamart, Habitat Ecologique

et Liberte

des Primates, Congo and BenoTt Goossens, Cardiff School

of

Biosciences, Cardiff University

Box 4.6 Julia Lloyd, Jane Goodall Institute-Uganda and

Lilly

Ajarova,

Uganda

Wildlife Authority

81

World Atlas

of Great Apes and their Conservation

David W. Liggett Iwww. da veliggett.com

82

BoNOBO [Pan paniscus]

Chapter 5

Bonobo
[Pan paniscus]
Carmen Lacambra, Jo Thompson, Takeshi

Furuichi.

HiLDE VERVAECKE, AND JEROEN STEVENS

bonobo {Pan paniscus Schwarz,

The

known as

also

pygmy' or

the

chimpanzee, occurs only


Basin

Central Africa, where

in

chimpanze nain or chimpanze

Bonobos have black


legs,

the inner

the Democratic Republic of the

of

(DRCI,

in

and

Congo

tail.

long as their

Perhaps the most

obvious physical differences from chimpanzees are


visible in the

adult

head and

the

(conventionally the

and

range

forest,

rence

tinctively

parted

in

The Congo River and the mountains


Albertine Rift isolate the bonobo from

bodied primates
the species

was

all

all

other large-

bonobo throughout

modern range has been well documented


via

other

not scientifically described until

1929, the existence of the

1880s

of the

baboons). Although

(including

explorers' journals,

its

since

naturalists'

photographs, missionaries' reports, and colonial


administrative records.'^
Field surveys of

"

proved very

difficult to

date.''

in

' "

the early
'^-

It

has

obtain a clear view of either

the total geographical area occupied by bonobos,

or the

likely

number

of

individuals within

(estimates range from 10 000 to over 100 0001.'^

it

the

in

dominated by moist

patchiness of the inner Congo Basin, combined

disturbance and

(to

the

forest,

in

swamp

grassland and dry forest.

of

far less than their

is

may

of

their area of actual occur-

maximum

geographical

be less than 30 percent

may be as low

of

as 0.25

it.

indi-

viduals per square kilometer.'^ The dotted range

polygon shown

in

tvlap

and based on 2004

5.1

distribution data represents an area of 373 585 km'.

The

total

bonobo population, assuming

a 30 percent

occupation of this range at a density


is

assumptions about

parameters.
in

of

0.25/km^

makes

28 019 animals. This indicative figure

significant

is

It

possible to place

the estimate^^ that


(in

in

the northern

Research), llongo

(in

Lomako, Lilungu, and

the

all

relevant

more confidence

the well studied sites of

Luo Reserve

for Scientific

the southern Luo Reserve),


Yalosidi, a total of

4 421 bono-

bos occurred, although the Yalosidi population


has now been
sites

*^

servational environment, and inherent ecological

human

bank] of the Congo River

between 300 and 750

lying,
is

and grassland,

range and

(at

lost.

Recorded densities

range from 0.35/km*

(at

at

these

five

llongol to 3.46/km'

Lomako).^^

Bonobos have been recorded in the Lomako,


Wamba, llongo, Lomami-Lualaba,

This uncertainty reflects the scale, challenging ob-

with the impacts of

and

and mosaics

Wamba

bonobos began

1970s and have continued to

'left'

Population densities

the middle.

apes (including chimpanzees], and

in

Although bonobos occupy a mosaic habitat

and hardly any


long and dis-

low

is

elevation,

forest

is

limited

is

major tributary the Lualaba; and,

its

bonobos have a short and rounded skull with

beard. The hair on their heads

bonobos

of

and west by the south bank

north,

east,

face. Generally speaking,

a black face, red lips, sidewhiskers.

the

DISTRIBUTION
The geographic range

south, by the Kasai/Sankuru river system. Their

noir.'''''"'

arms as

fur,

no

a tailtuft but

Congo

known as

is

it

19291,

graclle'

Kokolopori,

Salonga, and Lukuru regions, and


the west of Lake Tumba."-

be absent or

low density

small areas

in

' '" ""

the central parts of

south! the complex gradation between dry forest

to

and savanna.

the Salonga National Park, and absent from

at

in

to

Bonobos seem

much

83

World Atlas

Map

Bi

5.1

of Great Apes and their Conservation

Bonobo

distribution

Da'.i ~-n,,rrrs

are provided at thp pnd of

this

chapter

BoNOBO [Pan paniscus]

Mal-Ndombe and Salonga.

area between Lac

of the

The other area

absence

of

between the

falls

Tshuapa and Lomami Rivers (Map

bonobos have been observed


for a share."

are

5.1],

more

beg the meat holder

to

has been suggested that bonobos

It

able to source dietary protein from non-

reproductive plant parts than are chimpanzees, so

may have

BEHAVIOR AND ECOLOGY


studies of bonobos

tilost

carried out
of

the wild have been

in

primary forest

in

less

need

Bonobos

Habitat and diet

the northern part

in

but recent worl< has con-

their distribution,

for food

Lukuru

at

was observed

at Yalosidi

in

secondary forests. High densities are found, es-

stems

in

patchwork and edge-

'^

habitat mixtures.''

forest,

studied

in

the

Wamba

region."

rainy

A tendency

season

is

where they pre-

dry forest,

They also use

swamp and

tion of protein-rich food that

round.

large part of their

which apparently contain

forests,

dry forest,

to

and disturbed forest have been

home range comprises


fer to sleep.

lAllsmataceael, Pycreus vanderysti (Cyperaceael,

on the

recognize, but

When moving

in

trees,

they employ a method that can be described as

'quadrupedal scrambling', an inefficient

been seen

to

means

trees

in

is

sitting,

of

be sustained

over more than about AO m. The most


posture

common

(ZIngiberaceael.

spp.

Cyclosorus

included

Other

dentatus

Panicum brevifolium (Poaceael,

(Thelypterldaceael,

(both Marantaceael, as well as

Gambeya

" Bonobos have

(Sapotaceae|.'

also

lacourtiana

been seen

streams, and scooping up

In

handfuls of dead leaves, probably


"
brates and small fish.'

to

catch inverte-

Although bonobo diets are generally similar


across their range, differences have been noted

between populations studied


time.'

'

The extent

periods of

for long

which such differences are

to

rather than dependent upon

cultural

availability

is

resource

unclear

Social behavior

Bonobo groups number between


animals" and travel around 2
day,'' foraging

mainly for

cludes leaves,

pith,

mushrooms, and

km

10 and 120

on average each

Their diet also

fruit.

In-

flowers, seeds, nuts, shoots,

algae. Additional food

sources

include high-quality terrestrial herbaceous vege-

earthworms, larvae, termites, ants, honey,

truffles,

eaten

species

and

for both feeding

resting.'""

tation,

Aframomum

and

usually lead either to the base of trees from which

that has not

consumed throughout

most common were Ranaiisma humile

congensis, and Sarcophrynium schwelnfurthianum

of their traveling

they teed, or to nesting sites.

amphibious herbs and

the species

slapping the water

difficult to

marsh
on the

higher propor-

noted.

Bonobos do most

feed

to

Renealmia africana (ZIngiberaceael, Marantochloa

available all year

is

visit

disturbed

use drier habitats during the

to

ground. Their trails are

movement

Among

the year, the

Bonobo groups with access

swamp

of certain aquatic or

grasses."*

One group

pools.^"

grassland within the rain forest

and

Wamba

streams or

In

frequently to

firmed that they also use open savanna and

pecially in the latter,

hunting.'^

In

Lomako, and

have been seen foraging

marshlands, and

energy

to Invest

Yalosldl,

at

and aquatic

Subgroups or

Bonobo
fission

'parties' are

or group.

comes

to

30

from two

largely

sloned

site,

of

is

characterized

by

small temporary subgroups

Our knowledge

Equateur province

more
of

stable,

community

bonobo social systems

field

sites,

both

in

the

DRC: Lomako, an unprovi-

and Wamba, where provisioning used

occur Researchers provided food

to

formed by two

of

(parties! within a larger,

''^'

plants.'

organization

social

and fusion

bonobo groups

at

Wamba

to

some

prior to the cessation of

together with parties from other groups.''-*''^ As a

when research began again after


was not resumed.'"
The bonobos at the two sites show a number

com-

of similarities in social organization, but differences

individuals

and are composed

males, females,

of

and their offspring. They do not usually forage

presumed adaptation
petition In

to the greater feeding

small patches, the size

of

feeding parties

may vary with patch size." Bonobos occasionally


Including flying
consume small mammals," "^''
''^

squirrels. Infant duikers,

and

bats.

There

is

little

information on their hunting methods, and hunting

does not seem


is

available.

It

to
Is

be a frequent practice.

When meat

treated as a valuable resource;

studies

In

the

war, this provisioning

civil

1996;

have also been noted. As

community

Is

in

chimpanzees, the

the largest mixed-sex social unit of

individuals that maintain a closed social network.

members share a discrete,


home range; extensive overlap
between communities may occur and there may be
seasonal and yearly variations in home ranges."'

single community's

relatively

large,

85

World Atlas

of Great Apes and their Conservation

of

such subgroups requires prolonged observation

and detailed analysis

abundant data on individual

of

associations. Their occurrence

obvious

in

As

the

not necessarily

is

field.

chimpanzees, maturing males tend

in

remain within their natal

to

community, while

(birth)

maturing females leave and move from community


to

community before

result

IS

that both

made up

munities are

down

settling

to breed.

The

chimpanzee and bonobo com-

males that are more

females and

of unrelated

be related

likely to

one

to

number of difmuch longer time

masks

another. This similarity

ferences, however, such as the

period (years rather than months] during which

young female bonobos move between groups before


settling

down, and the very strong bonds that

Bonobo

structure

social

exist

"

between bonobo mothers and sons.^'"

dominated by

is

female coalitions that influence mating strategies

and food allocation. Females are smaller than


males, but maintain their social status through
cooperation with each other Female bonobos are
very skilful
Frances White

Grooming between an

Communities contain between 10 and 22

adutt male, adutt

female, and her infant,

Lomako and between 30 and 120 individuals in


Wamba." Tliere are almost equal numbers of adult

from the Bakumba

males and females

community

one Lomako community, the adult sex

Lomako

of the

forest.

individuals

in

in

Wamba,*' whereas

strongly female biased.

much

often

less

bonobo
terms

ratio

is

^''

daily

" The smallest functional


life

is

remain

in

fission

and

change

to varying

fusion,

in

males

of

parties

can

death

birth or

of

members;^'

females than males.'

Wamba,

in

with an

Lomako, parties

in

8.

37,

i8,

91

bonobos contains subgroups

others.

of individual

their

sometimes displace

preferred feeding

into
is

related to age,

males relates

positions.

whereas the status

to that of their

more

A community

of

prime have been observed

the death of their

mother^'

is

strual cycle and

home

share specific

range. Discernment

is

bonobo society

in

independent

maintained

munity, there are on average


interested

in

much more

to

is

polize

in

is

impossible.'''^'

in

these circumstances,

difficult for a

bonobo com-

chimpanzee
it

would be

high-status male to

mating opportunities, so male status

important to

individuals."^^^'

men-

many more females

mating than there are


In

of their

pregnancy and

into

a result, at any given time

form parties with each other than with

These subgroups tend

females

of

lactation, while further conception

community.

of

in

to fall in status after

thought to be related to their sustained sexual

attractiveness. This

As

to rise in

mother

The high status

individuals that

parts of the community's

86

get

to

Female status

members,

between one and 16." Parties usually

often

give charging displays

they are excited, females

status through the support of their mother; males

of

contain mature individuals of both sexes, with

more

may

bonobos

to that of adult

membership

contain about five individuals on average, with a

range

that adult female

is

Young adult males have been observed

stable parties are seen

13

result

have a social status roughly equal


males. Though males

degrees over days, hours, or even

to

males when necessary.

when

or by their permanent intergroup transfer Larger,

of

stra-

often than with males, interacting sexually

of

minutes. By contrast, membership of communities

more

The

and forage together^'Through

changes only with the

average

more

at

proximity to one another, or within earshot of each


other,'^ or that travel

"'

with other females, and forming alliances against

are

sustained

"

"

desirable food, sharing food with other females

unit

the party, defined variously

of the individuals that

establishing and maintaining strong

employed include controlling access

The net

Lomako than they

at

tegies

'-^'

communities are observed together

Entire

Wamba.'"-

least

in at

in

bonds with unrelated females."

monois

less

Male bonobos are rarely

BoNOBO [Pan paniscus]

observed

Males

to

compete or figtit over access

freely

access

females, and treat

them

counters are rare

proceptive

bonobo groups sometimes come together

manner;

a friendly

in

to females.

and

receptive

is

it

female that determines w/hether copulation

the

occurs. There

is

some

evidence, however, that high-

more success

ranking males have

suggesting that competition

lil<ely

to

feed

to

peaceful atmosphere. Male bonobos

or rest

in

become

excited and tend to stay behind the line of

contact

between the groups during such en-

counters, but females willingly enter a different

Compared

is

male chimpanzees, male bono-

to

have sired any particular offspring. This lack


over paternity

clarity

of

whether he

tell

In contrast, different

group, and will copulate with unfamiliar males."

not entirely absent.

is

system, no male can

In this

""

mating,^'

in

bonobos.

in

consistent with the

is

observed generalized paternalism: adult male

bonobos are extremely caring and affectionate with


sharing both food and

infants,

nesting

Amongst macaques, paternalism

spaces.

known

is

Box 5.1 SEED DISPERSAL BY BONOBOS AND THE SURVIVAL


OF RAIN FOREST

to

be

As specialized

frugivores,

bonobos are essential

long-term survival

for the

Lomako

of

associated with promiscuous female sexuality,

the rain forests

single-mount ejaculation and an even intragroup sex

together with seven other primate species, but they are the only one to ingest

ratio;'^

it

seems

bonobos.

in

that

something very similar occurs

Relatively food-poor environments,

seem to give rise


system among macaques,
however,

to the opposite social

ual encounters

between the same

a lack of paternalism.

The bonobos relaxed social

and sexual system has therefore been attributed


their diverse diet

to

Bonobo females indulge

bonobos occur

forest,

Bonobos are

liana.

number

excellent seed dispersers for a

more

they are primarily frugivorous |up to 70 percent or

First,
IS

and

ripe fruitl

rarely

damage

of the diet

the seeds consumed.

Second, they are large bodied and have simple guts, so that even large

seeds can be swallowed whole and passed undigested


seeds

some

of

fruit

bonobos

Third,

among

species are very large,

Anonidium mann/7 lAnnonaceael. These

collaborative

in

and

over 50 seeds that each measure 3

and resource-rich environment.

the

In

live.

of reasons.

before

pair

ejaculation happens), near-certain paternity, and

which they

regularly and disperse the whole seeds of a wide variety of species of rain
forest tree

harem-like sexual

of

multiple-mount ejaculation (repeated sex-

control,

in

cm

fruit
in

via

the feces.

weigh 3 kg

lor

morel and contain

length and weigh about 10

travel long distances

The

the biggest being those of

g.

and maintain large core areas.

genital rubbing, genital-genital contact,

and num-

Individuals cover over half their

erous related behaviors. The bonobo

clitoris

than 90 percent of their time within primary ram forest, thus providing long-

large and shifted ventrally

compared

chimpanzee."^' Genital contact


of

bonobo

social interaction, but

is

to that of the

common

is a

when

monopolized by an

individual.

that sexual activity

among females may

Hence,

it

relief

food

is

serve to
of

coalition

contact

is

relationships.

The frequency

of

genital

also related to female status, with low-

ranking females initiating contact

more

often than

high-ranking ones.^' Together with the lack of restriction in

heterosexual mating

more. The seeds


Fifth,

away

activity, this suite of

and

The high social status

in

of

bonobo

sexuality.

females may also

be related to the difference between chimpanzees

and bonobos

in

intergroup relationships.

Among

bonobos, intergroup interactions are frequent and


are

characterized

rather than

conflict.^'

by

excitement

Chimpanzees are known

antagonistic towards and


of

high-pitched

sometimes

kill

to

be

members

and spend more

on them

rely

TrecuUa afncana (Moaceael that weigh 10 kg or

later be dispersed

even further before being defecated.

bonobos do not sleep where they have been feeding, but move

to build nests

fruits that

and sleep

for

weigh about

in

trees elsewhere.

and

of tree

appear

liana

dispersal.

have evolved with bonobos

to

Carpodinus

lApocynaceael

gentilii

cm

kg with a hard rind, 2.5

thick, that

has

smaller-

bodied monkeys are unable to open. Others, such as Pancovia laurentii


(Sapindaceael, have seeds that germinate readily after passing through a

bonobo

gut, but not at all

if

the fruits

fall

uneaten beneath the parent tree or

are artificially planted, even at a distance from

sexual behaviors has led to an unusual amount of

research and popular interest

fruits of

may

Many species

social

tension, thereby serving to restore and maintain

year,

Fourth, they often carry fruits long distances before sharing and eating

them, such as the

thought

is

community range each

distance dispersal within a suitable habitat for rain-forest trees.

part

more frequent

after an episode of aggression or

promote reconciliation and the

is

The

disperser of

perser of

many

some

them. Of 130

bonobos have been seen

study,

fruit

that

it

is

the

most important

DRC, and may be the only

dis-

species collected and measured

in a

rain forest tree species in

of

it.

bonobo means

dietary diversity of the

to eat 63.

The

list

of fruit

species

be eaten by bonobos increases with each year of investigation," so


likely that

trees

in

in this

bonobos are involved

the inner

in

the dispersal of half or

Congo Basin. Without bonobos,

ecosystem would be

likely to

more

therefore,

known
it

to

seems

of all fruiting

major changes

occur within very few generations.

Frances White

other groups, while aggressive intergroup en-

87

World Atlas

of Great Apes and their Conservation

It

is

known why bonobo

not

social interactions

are so different from those of the chimpanzee. The

general view

that the high level of

is

bonobo female

sexuality associated with their being receptive for

extended times relieves a chief cause

male

through the abundance

friction

opportunities. '" "

male-

of

mating

of

The use by females

themselves may result

among

sexually egalitarian

in

their

of

sexuality for maintaining effective coalitions

which male possessiveness would

society, within

be ineffective, even

if

attempted. Frequent groom-

between the sexes reinforces social bonds and

ing

contributes to a relaxed social system.

Nonreproductive sexual behavior,


social interaction,

potentially costly in

is

like

all

terms

of

energy expenditure and reduced foraging time, so

it

cannot be ruled out that a food-rich environment

is

necessary enabling factor for the bonobo social

system. On the other hand, while there

Bonobo
nursed

more sexual

infants are
until

much

is

bonobo group than

chimpanzee one, among bonobos "instead

they are five

in

of

an

we see a social life peppered by brief


moments of sexual activity,"" so the energetic costs
may not be very great.

endless orgy,

years old.
David W. Liggelt Iwww.daveliggett

bos participate

in

less physical competition for copu-

lation opportunities

and interact less aggressively

with males of other groups. They do not participate


in

raids on neighboring communities.'""

philopatric social organization

species,

in

common

" The male


to the

two

which young males stay with the natal

group and young females migrate, clearly does not


predict these

aspects

distinctive

their

of

behavior More generally, bonobos

of

show much more frequent and

varied

evenly dispersed between individuals

social

both sexes

behavior than chimpanzees." Grooming

sexual
is

more

among bon-

Development and reproduction

between

individuals of the opposite sex

is

more

and

Details of reproductive development,

chimpanzees and bonobos

cycle, are given for both


in

Chapter

occurs
nine,

at

and

the

brief:

In

3.

seven years

adult size

full

first

of age,

genital swelling

sexual maturity at

reached

is

at 16. At eight

years of age, young females start to move between


groups; settling

and

The
is

in

new group occurs between


when

offspring are born

first

between 13 and

15.

the female

Only one infant

produced per pregnancy, often during a

grooming between females or males only^'

from March

It

has been proposed that the differences


intrinsic

than

nine

13.'"

frequent and occurs for longer periods of time than

between the two species may be less

of hor-

monal and behavioral events during the menstrual

obo than chimpanzee communities, and grooming

to

The menstrual

May during

usually

is

birth

peak

the light rainy season."

cycle lasts 36-46 days,

and the

gestation period has been estimated at 220-230

had been believed, and could be explained partly by

days. Infants are nursed until they are five years old,

environmental conditions including food supply and

distribution, party size, and sexual opportunity.^

and the mean

According

to

this view,

reduced competition be-

tween females enables more stable parties

to

be maintained, with more female sociability than

occurs

in

chimpanzees.

Some

scientists

also

consider that the genetic relationship between

bonobos and chimpanzees may be closer than the


evidence from comparison

DNA suggests.""

88

activity in a

of their

mitochondrial

birth interval

is

i.6 years.

It

is

not clear

whether menopause ever happens, as continued


menstrual cycling has been observed
are 45 or

more years

old."

The

in

females that

typical

tancy of between 50 and 55 years leads

age

of

five

life

to

expec-

an aver-

or six young being produced during

a lifetime."-''

Observations during 1976-1996

in

the

Wamba

region concluded that bonobos there have an infant


mortality rate that

is

much lower

than

is

recorded

BoNOBO [Pan paniscus]

for

chimpanzees. This

some combination
baceous foods

at

of

is

thought

Wamba,

from

result

to

the abundant

fruits

and her-

larger food patch size,

different locations

have been noted

to

preferences in the type and location


"^

have different

of trees

chosen

for nesting.

male

better female access to prime feeding sites,

Interactions with other animals

paternalism, and absence of infanticide."

Various diurnal species of

monkey share

the bon-

swamp monkey

Vocal behavior

obo range, including the Allen's

Bonobos are much more vocal than chimpanzees

[AUenopithecus

or any other great ape (see Box 5.21. They use

ILophocebus aterrimus], golden-bellied mangabey

numerous
distances,

calls that are audible over long

comat all

monkey

day and night. Hooting occurs most

guereza

monest long-distance

in

redtailed

and can be heard

when

echoes.

like

noticeable are the

frequently

(Cercocebus gateritus chrysogaster],

mangabey

high-hoots' that are the

synchronized choruses that

including

of the

black

monkey ICercopithecus ascanius], Congo Basin


Wolf's monkey [Cercoplthecus pogonias wolfi\, De
Brazza's monkey [Cercopithecus neglectus], dryad

end up sounding

times

and short

nigroviridis],

call

Among

the most

the bonobos arrive at feeding sites

the early morning and while they occupy a

prospective nesting site

in

the afternoon.

Other

or Salonga

or

guenon [Cercopithecus

black-and-white

colobus

dryas],

[Cotobus

guereza], and Thollon's red colobus [Procoiobus sp.


thoUoni]. At Yalosidi,

there

was

it

was concluded

dietary overlap

that although

between the bonobo and

vocal sounds have been identified during feeding

and copulation, and

response

in

to

Bonobos

danger^

in captivity

commonly make use

Little tool

of

implements, suggesting

Tool use

use by bonobos has been observed

wild. In the northern sector of the

in

the

Park, bonobos have been seen digging with sticks

in

termite mounds,'^ and males have used branches

in

displays. In captivity,

toot use in the wild.

Salonga National

bonobos use various objects:

rope swinging; self-wiping with leaves; and using


sticks

as

ladders or weapons

have

been

all

Captive juvenile bonobos have also

reported."

been seen using leaves

in play,

covering their eyes

and feeling their way around while blindfolded.'"


likely that

some

of the

same behaviors

It

are found

is

in

wild bonobos.

Nest building
Both day and night nests are

built

by bonobos; they

are used for sleeping, grooming, feeding, and playing.


Built afresh

every day, the nests are circular

shape and can measure up


Night nests are the
to construct.

to 1.3

materials from up to six trees

built in

the middle

in

may

gather

the construction of

whereas chimpanzees

use the branches and foliage

of only

purpose. There are also reports

ordinarily

one tree

of

for this

ground nests,

possibly built for purposes other than resting.

Females

in

diameter

above the ground], while day

nests are usually higher up. Bonobos

their night nests,

in

more elaborate and take longer

They are usually

canopy 115-30

build

'''"

higher nests, do so more

frequently during the day, and use

periods of time than

do males.

them

for longer

Bonobos from

89

World Atus

of Great Apes and their Conservation

Box

BONOBO COMMUNICATION

5.2

levels of

symbolic communication.

for

If,

any reason,

change

the decision about theirdestination needs to

Bonobo communication
great apes

number

in a

appear

which)

to

from

differs

ways,

of interesting

be related

of otfner

tfiat

have adopted social strategies unlike those

Bonobos are

apes.

munities

up

of

to

often found

in

of

other

large stable

com-

20 individuals that

move and

together, and break up into smaller parties

come

they

to the

contrast with

ground

sound

their

feed

when

for long-distance travel.

By

other great apes, bonobos are

all

extremely vocal, both

where

all of

bonobos

to ttie fact that

captivity

in

and

the wild,

in

limited mainly to the

is

Bonobos also engage

canopy

freguent exchanges of

in

glances and gestures, drag branches, ostentatiously break multiple branches,

and pound on tree

example,

floor (for

way
to

to a

feeding

if

the forest

to

they encounter traps on the

site), this

must be conveyed

quietly

one another

Much more needs

be understood about

to

bonobo communication systems

the wild. Like

in

other great apes, bonobos spontaneously begin to

human speech and

understand spoken

written symbols with that speech


(see Box 3.1

These competencies require neither

1.

nor rewards. They emerge

training

when communicative

especially

pair

to

captive settings

in

bedded within the


place to place

intuitively,

em-

activities are

daily activity of traveling

from

the forest to locate food resources.

in

buttresses as acts of communication. Preliminary

Captive bonobos as young as two years of age can

observations suggest that they are able

easily mentally

messages using crushed vegetation

to

to

leave

indicate

The need
two

coordinate group travel between

to

to four fruiting

level

sources per day requires high-

communication.

community

not possible for a large

is

to

and arrive

sufficient

find

randomly about the

traveling

locate

It

bonobos

of

made

is

for the

fruit

must agree and end up


hours

community

at the

same

food by

They must

forest.

ripening

at

resources

When

throughout their environment each day


decision

to travel, all

location several

even though they neither see each

later,

other nor vocalize as they travel on the forest floor

Because bonobo communities are large but


travel on the

ground

in

small quiet parties, each

out of sight of the other, their daily lives require high

the four

were

all

monkey species found there, the monkeys


much more arboreal than the bonobo, and

their ecological niches

were narrower."

species of primate have been observed,


ticular

with

Angolan

the

colobus

even guide
to

in

par-

IColobus

intentions,

treatment

killed the

were not seen


result in

to

harm

encounters, rough

monkeys. The dead monkeys

be eaten. Interactions that did not

to

any

including grooming

of age,

of the individuals involved,

in

Wamba."

of age,

their travel

destinations

in

they can decide

of travel.

By eight

where others should

so inform them, and then wait for their

travel,

Local trackers

return.

bonobos send scouts

DRC

in

to

has yet

to

that

report

wild

check out nearby food

resources as they travel from point A


activity

be verified

among

to B.

While this

wild bonobos,

it

does appear among captive bonobos.

POPULATION
Status and trends
population trend information

is

available for

bonobos, and the impacts of the war are so far largely

unknown. The loss


research
is

of the population at the

site of Yalosidi

former

has been confirmed, and

thought that numbers have declined


site.

The

total population is

much reduced because


ularly

the

spread

of

of

human

thought

at the
to

be

activities, partic-

firearms (including,

recently, powerful military

more

weapons], together with

habitat clearance.

between young bonobos and

red colobus {Procoiobus badius], have also been

observed

means, and

to their forest

disagree with each other

to

about their intended destinations


years

it

of the

two or three

plan

advance, and begin

Wamba

some

human companions new

bonobos can answer questions about

mostly treated these monkeys as they did their


but, in

forest, travel to food

designations the bonobos select and specify

angolensis] and redtailed monkey." The bonobos

own species

ha

a 20

through symbolic means." By four years

Little

between bonobos and other

Interactions

map

resources by previous or novel routes, communicate


their travel intentions through symbolic

direction of travel.'"

90

come down

once the bonobos have

The patchy

distribution of the species hin-

ders the estimation


their

known range,

of

population numbers. Within

local population densities

range

BoNOBO [Pan PANiscus]

bonobos

Captive

acquire

keyboard Isee Box

lexigram

via

can

productive

hundred words, expressed

of several

vocabularies

and are

3.11,

comprehend several thousand

believed to be able to

spoken words. Bonobos combine symbols without

grammar

being taught to do so and use a simple


that

own

partially of their

is

construction. Their

tures far exceeds what they produce with lexical

may be an

symbols, but this

artifact of the

symbol boards rather than a


grammatical

their

symbolic dialogs that

and span several

way

to another,

ability.

may

ing the core

left

later point in time.

They have no
to another,

turn giving

in

to the dialog at a
difficulty in leaving

to

develop this

of

in

skill

immediately adopted the developed form without


the intermediate stages, after observing the

first

bonobo."
Tools, language,
in

reared

and culture appear to develop

manner among

a coordinated

dialogs,

captive

bonobos

an appropriate environment; the same

in

be true of wild bonobos. Vocalizations

wild are complex, frequent,

in

hammer

hand and the

Bonobos engage

run for 20 or 30 minutes

and may return

moving

the

The second bonobo

the right

likely to

one

topic,

in

developing the use

finally

'throw-like' blows, while hold-

reflection of a limit

each topic

topics,

unnatural

downward

glancing

understand complex grammatical struc-

ability to

to

and throwing, before

in

is

the

exchanged as lengthy

and accompanied by pointing gestures.

may occur

Distinct cultures

at different sites,

significant discoveries about

and

bonobo culture and

communication doubtless remain

to

be made.

Susan Savage-Rumbaugh

and then picking up

the former topic, without any need to recreate the

former conversations that


Their
setting

ability to

up

led

A young bonobo

to it.'^"

acquire language

in a

hooting.

captive

David

Liggett Iwww.daveliggett

V\/

comi

paralleled by a similar capacity to acquire

is

the rudiments of stone-tool manufacture." While

symbolic language and stone-tool manu-

both

facture

were

initially

demonstrated by human

companions, the bonobos acquired these

abilities

through skilled imitation and observation. Even

more

intriguing:

these

skills,

once one bonobo had acquired

they were transferred to other adult

bonobos and

without the need

to their offspring,

human modeling,

for

with

often

far

greater

efficiency than that associated with the initial skill

For example, the

acquisition.

bonobo

first

to

acquire the techniques for stone-tool manufacturing

went through phases

from 0.25

to

3.7/km^ but the species

'"-

common."-

as Endangered

''''

in

"'"''

Red

nowhere

is

The bonobo

the 2004

knapping

of horizontal

classified

is

Lisf of

lUCN-The

World Conservation Union, indicating that


very high risk of extinction
future.
of

the

The bonobo

Is

Convention

Endangered Species
(CITES), which

DRC

in

the wild

also Included

on
of

joined

in

In

International

Wild
In

it

has a

the near

Appendix

Trade

in

Fauna and Flora

1976.

Is

important to the conservation

A systematic line-transect and reconnaissance survey was completed across about


of

61

bonobos.

percent of the 36 560 km^ park

the Congolese

in

2004, under

the fvlonitoring of the Illegal Killing of Elephants

of

CITES." This survey was co-

States

Fish

for

Institute

(ICCNl and funded by

and

Nature Conservation

WWF International, the

Wildlife

the

Service,

Research Project. The Lukuru


Project and the
fieldwork.

patchily; in

Max Planck

Wildlife

Institute

Bonobos were found

some

sectors,

Relatively high densities

to

United

European

Community, USAID-CARPE, and the Lukuru

in

As the only national park within the bonobo


range, Salonga

IMIKEI program

ordinated by the Wildlife Conservation Society and

Wildlife

Research

were involved
be distributed

none were encountered.

were found

In

parts of both

blocks of the park: the north and northwest of the

northern

block

and the southeast, west, and

northwest

of the

southern block, as well as

In

the

corridor that separates the two blocks.

91

World Atlas

of Great Apes and their Conservation

and rudimentary healthcare provision, human

Threats

Bonobos were probably always


in

tlie

patchily distributed

diseases or parasites

Congo

bonobos."

vast area of the inner

Basin, and

hunnan pressure and forest fragmentation have


reduced their distribution," while

meat has eroded

economy as

agricultural

the

Lomako

forest." Although

quite well in

secondary

identified as the

The

latter

hunting of bonobos

of the

and com-

areas such as

in

bonobos seem

do

to

has been

forests, logging

most important long-term threat

to

is

National Park, Luo Reserve for Scientific Research,


Kokolopori,

700 km." There was

logging are difficult to disentangle from the sim-

tive

hunting, as both access to

in

demands from

markets

local

Other threats

accelerate.

population

for

bonobo

to

and can

in Africa. In

1999, there

expected
life

in

response

to

DRC

population
shelter,
IS

to

in

were 60

is

double within 25 years." Conditions

of

still

and

Pressure on

more

much

all forest

of

the

resources

Where access along

rivers

is

human immigration and land-use change

an order from a hawker

is

(middle rnanl from

taboos, an increase

Kinshasa.

Where many people

frequent.

In
in

the absence of protective

hunting pressure

settle with only

Jo Thompson/Lukuru

is

likely.

poor sanitation

V^ildlife

injure or

Research Project

around 50 loads

in

of

- but snares are not selec-

bonobos that encounter

kill

them. Elephants, by contrast, are targeted for their

meat by commercial hunters using semiautomatic


weapons, and are viewed as under severe

mainly

supply

to

threat.

bonobos,

of trade in live

Where

collections."

private

hunting does occur, females with young are par-

as a threatened mother

ticularly vulnerable,

will

when it has grown to half


slows her down and makes her an

carry her offspring even

her

size.^

This

easy target.

on forest products for food,

relies

fuel.

increasing rapidly

possible,

million

number

are very difficult and

'

There are also reports

increasing at

is

people; at current growth rates, this

hunters

in

evidence of

direct

little

bushmeat examined

growth rate

by local

a few. In

bonobo meat was present

almost 3 percent per year, the highest annual

killed

name

with bonobo distributions." The direct impacts of

DRCs human

was

to

snares and 97 hunting camps were found

habitats include cultivation and mining.

it

Wamba, and Lukuru,

the 200^ reconnaissance survey at Salonga, 339

bonobo hunting - one skull was found, and no

bushmeat

carried

it

logging concessions - currently only partly active -

the forest and

from the forest after

a growing threat, although

is

so far thought to be absent from the Salonga

the species because of the widespread overlap of

ultaneous increase

A bonobo being

Congo Basin, bonobos are

much

a result of war,

mercial hunting has intensified

parts of the

hunted and eaten by local people. Commercial

their numbers.^"

some

also be transmitted to

bush-

killing for

threat has escalated with the loss of

In

may

Since the start of the

armed

conflict in 1996,

some bonobos have been killed


including some at the Luo Reserve

by

soldiers,

for Scientific

Research."' Conservation and research programs

have also been disrupted, jeopardizing ongoing


studies of bonobo

life

history

and evolution, as

well as conservation programs involving bonobo

communities. These
scaled
the

have generally been

activities

down rather than

war has penetrated

halted altogether"

Where

has

led to

the forest,

it

increased local reliance on wild products, including

bushmeat." Adult bonobos have been

killed for

their meat, with juveniles being sold as pets. For

the 12 infant

bonobos seen

over a

month period

five

troubles,

been

in

the Kinshasa market

at

the

height of the

60-120 bonobos are estimated

killed."

to

have

Most bonobo populations, however,

are thought to have been unaffected by the war,

due

to their

remoteness from the

conflict area."

HUMAN ATTITUDES AND TRADITIONS


History and tradition

Humans

are thought to have inhabited the Congo

Basin for at least 100 000 years.' The great migration


of

Bantu-speaking people from the area now known

as southeastern Nigeria began around 1000 BC.

92

BoNOBO [Pan PANiscus]

They dispersed across the forests and savannas of


Africa, including the current DRC." As a result,

DRCs

population

connposed

is

different ethnic groups,

uages

Bantu

of the

most

family.

more than 250

of

non-Bantu

d'Amenagement
of the

Forestier,

bonobo's range

lies

within areas designated as logging concessions.''

Modern

which speak lang-

of

d'inventaire

guns and steel wire

tools such as

The largest are the Luba,

Increase hunting efficiency; even though bonobos

BC and 200 AD,

might not be the target prey, they may be caught

Kongo, and Mongo.*'' Between 100


the

Permanent

approximately 24 percent

sometimes l<nown as

peoples

in

traps and snares." Although local

human popu-

pygmies' had been driven into the central Congo

lations are familiar with bonobos, people often fear

and vanished elsewhere through inter-

them.'^ People and bonobos frequently share the

Basin,

breeding, depopulation, and cultural dominance by

the Bantu peoples.

country
istered
of

In

the late 19th century, the

was colonized by Belgium and was adminfor many years as the personal possession

Belgium's King Leopold

density
ing

was low and

Prior to this,

II.

depended on farm-

livelihoods

and hunting, probably

human

rough balance with land

In

resources. With colonization, however,

and

forest

new

technologies and domesticated species were

Europeans conquered and


area, exploiting

using

Its

its

major

Independent

of

some extent settled the

to

inhabitants and resources, and

DRC became

rivers for transport.

Belgian rule

in

Warfare

areas

a threat to

human

of

populations to

These movements have

bonobos because people with no

taboos against bonobo consumption have moved


into

bonobo range areas.

CONSERVATION AND RESEARCH


Protected areas

The minimum area needed

support a viable

to

300-600 km^ the exact figure depending on the

bonobo population

the Lukuru area, an important area for bonobo

threat,

research and conservation." Throughout the rest

areas, both over 300

Bantu Mongo people

range, the

during the 1990s forced the

relative safety

of

become

DRC

In

movement

large-scale

population of bonobos has been estimated to be

1960.

The Bantu Ndegense people now dominate

the bonobo

be

to

serious agricultural pests.

the federation of four ethnic groups that occupy

of

and bonobos occa-

wild food,

of

gardens but are not considered

sionally raid

methods changed.

production

and

Introduced,

same sources

and other

density,

levels

km'

and types

of

Two protected

factors."'

local

area, cover parts of

In

the bonobo range: Salonga National Park and the

Research. The Lomami-

dominate." Wild animals have been hunted by local

Luo Reserve

people for generations." Traditional hunting tech-

Lualaba Forest Reserve also contains bonobos, and

niques for domestic consumption use bow-and-

there are

arrow and nets, but guns are also commonly used,

status.

especially since the beginning of the


In

as

there

descendants

bonobos

that

belief

younger brother

that lived in the forest;

in

humans
of a

'fallen brother'

again.

These

who

is

some southern

trying to

its

when

to

become

beliefs support local hunting

taboos. Elsewhere, and even increasingly

zones,

are

bonobo

Is killed

by-products: the brain

is

there

is

in

bonobos

taboo

demand

for

considered a delicacy;

in

1970 largely
still

covers an area of 36 560 km'

in

within the geographic distribution of the bonobo.

DRCs

Service

to

and

two blocks

separated by an unprotected corridor 40-A5


wide. The park

by

swamp
It

encompasses

forest,

of

river terraces

to hold

that,

with associated
forest

although the park

good numbers

of

bonobo

Lomako and Wamba, there are


numbers In the northeastern part."

to

Government involvement and


are poor

km

a low plateau covered

and high plateaus with dry

has been reported

significant

DRC can now be allocated


lease; some have been awarded

intact

almost equal size ISecteur Nord and Secteur Sud\

compared

In

the center of the

Is

does not appear

Recent events

In

The park

protect the bonobo.''

wash and strengthen babies."

2003 figures from

area

in

cover

to

to full protected

it

Lomami-Lualaba, suppor-

bonobo range area was created

strength on men; and crushed bones are used to

According

to

2003."

riverine forests,

on a 25-99 year

upgrade

The Salonga National Park

the ashes of the bones are thought to confer great

Timber concessions

to

by Mwinylhall, confirmed the presence there of

family of

such as Lukuru, bonobos are believed

represent a

human

of a

is

moves

An expedition

ted by the Wildlife Conservation Society and led

war

northern parts of the bonobos' range such

Wamba,

parts,

civil

for Scientific

in this

application of laws

area, however,"

and hunting

is

current threat.'^

The Luo Reserve

for

Scientific

Research

93

World Atlas

of Great Apes and their Conservation

Tumba

by staff of the

DRCs

Centre de Recherche en

Ecologie et Forestrie, but the status of this proposal


is

unclear

Conservation and research activities


Wild bonobos can be observed only

in

DRC, so

field

research on the species has been greatly affected


by political events
to

in this

country. Scientists

explore the areas biodiversity

in

began

1973; academic

research and conservation interests prospered


during the 1970s and 1980s, with the encourage-

ment

of the

The

relatively dry

Lukuru area

war

civil

most

in

field

research, due to the

overthrew President Mobutu

that

much

1996/1997, affecting

is

in

country over the

of the

following years.

important for bonobo

Most

research and
conservation.

in

(358 km')

section

is

!:

._

jkuru Wildlife Research Project

bonobo research

which

of

southern section

in

is

in

site,

Wamba

the northern

village,

and the

was

llongo village. This area

1973 as an ideal place

identified in

to

study boncbos;

the

research on bonobos has been done

field

Wamba

and Lomako areas.

It

hoped

is

that

studies from Lukuru, a mosaic of dry forest and

savanna,

behavioral ecology

will provide insights into

not available from forested research sites. Other

research sites have included Yalosidi, Lake Tumba,

and Lilungu. Bonobo research has tended

to

focus

since then, local people have been involved as field

on social behavior, being driven by comparison with

and conser-

chimpanzees and also by public and academic

assistants and laborers


vation

projects.

(Inuyama,

fieldworl<

in

The Primate Research

Japan),

Institute

Wamba Committee

the

for

interest

in

bonobo sexuality and female

coalition

building, concurrent with the growth of the femi-

movement

Bonobo Research, and the Centre de Recherche en

nist

Ecologie et Forestrie together run a project based at

behavioral ecology of bonobos has also received

Wamba

some

village. Agricultural

are the main threats

in

expansion and logging

returned

in

The Lomal<o

forest

people.

in

and tool use have been

gated mainly

captive

has been reported that there

is

a viable

In

identified.

The

MIKE

forest

pressure

elephant

been affected by

hunting." Although local people traditionally hunt


for
is

bushmeat, bonobos are protected by taboo. This

changing, however, as the agricultural economy

continues to decline, and the area

is

becoming more

accessible. Efforts to establish a national park

in

the

area have not yet been successful.

the

in

Lake Tumba and Kokolopori areas, but no

program,

with

and

ape

and sun/eyed

in

to build

range states for managing

populations.

Through

program, bonobo populations are being

this

Identified

parts of the elephant range that

have not been previously or recently researched."

Assessments
carried

out

of
in

bonobo populations have been

Lomako, Lukuru, and Salonga

National Park, and a conservation Infrastructure has

been

Recent studies have also been undertaken

recently.

by the Wildlife Conservation Society, aims


institutional capacity in

the south-central part of the forest, but

more

survey

technical and administrative coordination provided

population of bonobos living relatively free of hunting


in

Investi-

bonobos, with follow-on

fieldwork being undertaken

100 km' bounded by

that the northern population has

habitats they

cation, language,

has good forest cover, and contains few


It

of

Meanwhile, communi-

war, scientists

the north of the bonobo

of 3

and the range

attention,

occupy has been

2002.'"'

range occupies an area


rivers,

civil

during the 1970s and 1980s." The

much

the area. Although

research stopped during the

built at the latter

The ICCN
country's

Is

two

of

these

sites.*"

responsible for managing the

protected areas and

related

research.

protected area has been established here." The

Supported by the United Nations Educational,

Bososandja Community Forest

Scientific

to the

south

range area has also been proposed for


protection.'"

96

former President Mobutu. There was

hiatus

later a

protected

area

of the
official

was proposed

at

and Cultural Organization (UNESCO) and

the United Nations Foundation,

it

has worked

conserve bonobos and other great apes

In

to

protected

BoNOBO [Pan PANiscus]

areas during the conflict

been involved

also

in

DRC.'^

Many

national and

nongovernmental organizations have

international

conservation efforts. The

in

Zoological Society of Milwaukee started a project

was

the Salonga National Park. This fieldwork

in

suspended because

armed

of the

conflict, but the

US Agency
among those

Zoological Society of Milwaukee and the

Development are

International

tor

supporting
Artists,

awareness

public

activities

DRC.

in

educators, nongovernmental organizations,

and government

have been involved

officials

research and education, and

in

producing booklets

in

and magazines." Other key agencies include the

Wamba Committee

Bonobo Research and

for

Wamba;

the National Geographic Society for

Lukuru

Wildlife

Institute for the

Chapter

profile in

likely to

of 100

activities

can be found

the

in

research

these extraordinary primates

much

be

An updated
a

more

movement
sure
tural

is

maximum

of

taboos erode with the

people within their range. This pres-

growing as warfare has damaged the agricul-

economy and encouraged

and commerce

in

forest exploitation

bushmeat. As the war

an end, moreover,

it

is

is

brought

logging and mining will escalate dramatically.

hunting

to

reach

human

population, leading to increased

of all wildlife.

Set against

this,

however, bonobos do occur

forest,

which remain remote and

the mid-1990s. The latter compiles

each

identifies conservation pri-

It

and recommends actions

site,

status), regulation le.g. habitat

and education and

November

the

1999.

include

the

increasingly

constituency of interest
partly

in

participants

a
Jo

redemptive contrast
Thompson/Lukuru

Wildlife

to

threats and

identified

meeting

set

Proposed

assessment

of
of

of information; public

research

sites.

the

Lukuru Wildlife

meeting with

lyo

Booto

Alfonse, Grand Chef

de Groupement de
Isolu.

The Grand Chef

human

has directed his people

activities included raising

to collaborate with the

species

DRC; coordination

of

Research Project,

priorities

also gave attention to the need to improve

of the

lives, offering

the stereotype

of the

to

research and conservation. Recommendations

life.

international

Jo Thompson. Director

lUCN Conservation
a

strong

the bonobo. This arises

Research Project

targeted species-based conservation measures, but

parties;

place

from the perception that these primates

assess bonobo conservation status, during which

awareness

in

and management

have charming and peaceful social

training.

Breeding Specialist Group conducted

quality of

relatively

become overwhelming. Other conservation assets

in

range and population

for

in

areas such as Salonga National Park and the

processes before pressures on these bonobos

plan.'^

involving research (e.g. determination of the current

In

As

new transport routes will allow


new markets among the rapidly

areas are opened up,

were

bonobo-focused action

the previous 20 years.

protection),

to

feared that industrial-scale

effective conservation, education,

information on research sites and activities over

in

estimate

action plan for African primates," and

specific

both published

orities

is

scarce, with a total population


less than the

unpeopled. The opportunity exists to put

Conservation priorities

commu-

sites."'

000 individuals. They are increasingly hunted

Lomako

16.

of the

for food as local protective

Kotal region. Further infor-

Lui

mation on conservation

DRC

Congo Basin, but

expanding

and the Max Planck

of

most

from secure. They are widespread over the inner

bushmeat

Kokolopori;

for

The future
far

Lac Tumba; Vie

Initiative for

priorities for the future, including

nity participation at

Research Project for Lukuru; the

Bonobo Conservation
Sauvage

the

some

infrastructure development and continued

in

mammals

1997 to assess bonobos and other large

established

among

activities

project

the peoples of

among

in

protecting

bonobos.

various

bonobo populations; sharing


education; and reopening of

There were also calls

international attention to

for greater

DRC, stronger

efforts to

restore peace, and investment to strengthen and

areas.'

further

on bonobos was organized

in

Japan during July

maintain

protected

2003. which

workshop

reviewed research carried out

Wamba. Lomako, and Lukuru. The workshop

at

also

fz^M:^
95

World Atlas

of Great Apes and their Conservation

aggressive chimpanzee as an alternative model for

guarantee

hominid societies.

greatly amplify the impact of conservation

enthusiasm could be translated

This

public support

for

donor governments

Such arrangements would

meaof

nongovernmental organizations, researchers, and

many

species and

the

'adopt'

to

survival.

sures otherwise based mainly on the enthusiasm

into

long-term commitment by

its

local people.

FURTHER READING
Bermejo,

Sabater

M., Illera, G.,

records from Lilungu

Coxe,

S.,

Rosen,

P.,

Lanting,

F.

Institute,

mushrooms consumed
Phmatology }5

International Journal of

N., Miller, P., Seal, U. 11999)

Primate Research

de Waal,

11994) Animals and

PI, J.

llkelal. Zaire.

Bonobo Conservation Assessment.

Fruth,

B.

Anthropology UU

(A):

Some

Horn. A. 11980)

Tumba,
Kano,

T.

J.,

eds 12001)

All

Apes Great and Small,

Academic/Plenum Publishers, Boston, Dordrecht, London, Moscow, New


(2003) Culture

vol.

York.

bonobos? Between-species and within-species behavior Current

in

563-570.

observations on the ecology

Zaire. Folia Primatologica

21*:

of

the bonobo chimpanzee [Pan paniscus. Schwarz 1929) near Lake

145-169.

Pygmy Chimpanzee Behavior and

11992) The Last Ape:

Kortlandt, A. (1996)

Worl<stiop Report. Kyoto University

11997) Bonobo. University of California Press, Berkeley

African Apes. Kluwer


G.,

new

879-898.

Inuyama, Japan.

Galdlkas, B.M.F., Briggs, N.E., Sheeran, L.K., Shapiro, G.L., Goodall,

Hohmann,

by bonobos [Pan paniscus] -

|6|:

A survey

of

Ecology. Stanford University Press, Stanford.

the geographical range, habitats and conservation of the

pygmy chimpanzee [Pan

paniscus): an ecological perspective. Primate Conservation 16: 21-36.

McGrew, W.C., Marchant,

Sussman,

L.F.,

The

R.L.. ed. 11984)

Thompson,

J.,

Hohmann,

G.,

Nlshida,T., eds 119961 Great Ape Societies.

Pygmy Chimpanzee,

Furulchl,

T.,

Evolutionary Biology

Cambridge University Press, Cambridge, UK.

and Behavior Plenum

Press,

New

York.

eds 12003) Bonobo Workshop: Behaviour, Ecology and Conservation of Wild

Bonobos. Inuyama, Japan.

Thompson, J.A.M.

Taxonomy and Ecology

11997) The History,

Description of a Wild Population Living

Thompson-Handler,

N.,

in a

of the

Bonobo Pan paniscus ISchwarz,

19291, with a First

Forest/savanna /Mosaic Habitat. PhD dissertation, University

of Oxford.

Malenky, R.K., Relnartz, G.E. 11995) Action Plan for Pan paniscus: Report on Free-ranging

Populations and Proposals for their Preservation. Zoological Society

lUCN/SSC Primate

of

Milwaukee County

in

cooperation with the

Specialist Group, Milwaukee.

MAP DATA SOURCES


Map

5.1

Great apes data are based on the following source, with updates as cited

Chapter

Butynskl.T.M. (2001) Africa's great apes.


Stevens,

in

the

DRC

country profile

in

16:

E.F.,

Arluke,

A., eds.

In:

Beck, B.B., Stoinski,

T.S.,

Hutchins, M., Maple,

T.L.,

Norton,

B.,

Rowan,

A.,

Great Apes and Humans: The Ethics of Coexistence. Smithsonian Institution Press,

Washington, DC. pp. 3-56.


For protected area and other data, see Using the maps'.

ACKNOWLEDGMENTS
Thanks
to

to Colin

Groves [Australian National University] for valuable comments on the draft

Stephen Blake, John Hart, and colleagues

surveys, and to Els Cornelissen (Royal

(Wildlife

Museum

for Central Africa,

AUTHORS
Carmen Lacambra, UNEP World Conservation Monitoring Centre
Jo Thompson, Lukuru Wildlife Research Project
Takeshi Furuichi, Meiji-Gakuin University
Hilde Vervaecke, University of

Jeroen Stevens, University

96

of

Antwerp
Antwerp

Box

5.1

Frances White, University

Box

5.2

Susan Savage-Rumbaugh. Georgia State University

of

of this

chapter Thanks also

Conservation Society] for information about the MIKE bonobo

Oregon

Belgium)

for archeological advice.

Gorilla overview

Chapter 6

Gorilla

overview
Julian Caldecott and Sarah Ferriss

There
the

are two species of gorilla, separated

pleted causes

them

be

to

tends

known as
be

'silverbacks'.

somewhat

larger

from one another by the inner Congo Basin,

The eastern

that region of Central Africa to the south of

than the western. Diagnostic differences between

Congo River

that

inhabited by bonobos. Each

is

gorilla

to

the two species include:

Eastern lowland gorilla,

species has two subspecies:'^

the eastern gorilla [Gorilla tiennge/ Matschie,

1903)

divided

is

gorilla [G. b.

mountain
19031;

into

gorilla

gorilla

b.

[G.

eastern

lowland

a silverback male,

western, which has sleeker and grayer or

Democratic Republic of

western gorillas

the Congo.

browner
Elizabeth

hair; the

head hair

of

A Williamson

beringei Matschie,

and

the western gorilla


18471

the

grauerl Matschie, 19U| and the

the eastern has longer, blacker hair than the

divided

is

[G.

the

western lowland

gorilla Savage,

g.

Cross River

gorilla Savage,

[Gorilla

into

gorilla

[G.

18471 and the

diehli Matschie,

g.

19041.

Mitochondrial

DNA (mtDNAl

research suggests that

the lineage of the western gorilla diverged from that

eastern gorilla around 2 million years ago

of the

(myal." This does not necessarily indicate that the

populations had already separated. Tropical Africa

had a drier and cooler climate

menting the forests


of the

in

gorillas lived,

at this

which the

time," frag-

common

divergence already underway as a result

geographical range

The two

ancestor

and possibly accelerating a


of

the wide

of the species.

gorilla

species

have

numerous

similarities" and, until 2001, were recognized only

as

subspecies."

Both are large and sexually

dimorphic, with adult males weighing up

200 kg and adult females around half

to

that.

about

Both

species have broad chests and shoulders, large


heads, and hairless, shiny black faces.

In

both,

maturing males develop a silvering of the hair on


their

backs and sagittal crests, which when com-

97

World Atlas

of Great Apes and their Conservation

Mountain gorillas have been studied high

in

the three national parks of the Virungas (Virunga,

Volcanoes, and Mgahingal, and at lower elevation


in

the forest of the Bwindi Impenetrable National

Park.

In

the Virungas, the diet

overwhelmingly

is

dominated by herbaceous leaves and shoots Iwhich


are

abundant

the Virungas, but

Bwindi the diet

while at

there],

more

contains far

fruits (which are very

common

at Bwindi).

scarce

'" ^'

in

both

In

places, the diet reflects seasonal influences, and

the gorillas gorge on 'seasonal specials' such as

bamboo

shoots, or

consume

more

diverse diet

including herbaceous vegetation, bark, and twigs,

as availability allows.
Consistent with the low quality and poor

mountain gorillas

digestibility of their diet,

Virungas spend

much

of their daylight

the

in

time feeding,

and otherwise rest." Their groups occupy a small

move

area for a day or two and then


returning for several

seldom

on,

months while the vegetation

recovers from being trampled and harvested. As

An

infant

herbaceous vegetation

western

low dietary

is

abundant, widespread, and

quality, there

lowland gorilla seeks

of

comfort riding

defend

'piggyback' on an older

cases, mountain gorilla groups

gorilla at the Lefini

completely overlapping

rehabilitation site,

Bwindi are more mobile than

Congo.
Ian

Redmond/UNESCO

tends to have red tones, with the crest and

nape hair

males usually being

adult

of

Virungas."'"'
of

seasonal

(some lowland'

the western silverback's saddle of white hair

the

body color than the eastern


tends

to

more

silver-

be more clearly

more against the

dark hair

eastern

montane
in

home ranges

same

the precise selection varying

kinds of foods, with

according

to

their

absolute and relative availability Their larger body

enables gorillas

to

consume

somewhat

poorer-quality diet than that of other great apes.


Dietary choice

is

constrained by their stomachs,

which are simple and nonfermenting and so preclude eating too

many mature

leaves.

of

of

forest," but the size of their

lowland tropical forest

is

gorillas! in

the

of

Congo

13-17 km'

in

home range

unknown. Like mount-

ain gorillas, they travel less

in

montane than

in

made up

of

is

seeds, leaves, stems, and bark as well as

ants, termites,

the

the

Itombwe forests

Democratic Republic

(DRCI- Groups occupy

fruit,

much

year,^'
in

gorillas live at

some mountain

lower-altitude forests. Their diet

ECOLOGY
All gorillas eat

at

living

Eastern lowland gorillas have been studied

the Kahuzi-Biega, Maiko, and

it

Those

the Virungas; a

in

Bwindi, and the corresponding

jaw musculature; and

delineated and to stand out

98

fruit at

higher altitudes than

which

some

In

the Virungas have

This reflects the higher availability

than the western, suggesting a more powerful

often extends onto the thigh, and grades

size

'

at various altitudes

back's, in

in

ranges.'^

sagittal crest along the midline of the skull

into the

ecological need to

group at Bwindi may use up to 40 km' in a


compared with 5-11 km'' typically used

has a more developed

gorilla

is little

against other gorillas.

increased travel to obtain such preferred foods.

striking chestnut color;"

the eastern

home ranges

and other insects. Fruits are pre-

when these are scarce, eastern lowland


gorillas eat more herbaceous vegetation. Large
quantities of bamboo shoots are eaten seasonally.

ferred but,

Western

When

fruit is

gorilla diet also varies seasonally'

abundant,

it

may

constitute

"''^

most

of

the diet but at other times shoots, young leaves,

and bark are eaten instead. Terrestrial herbaceous


vegetation, aquatic herbs, and insects are eaten year

Gorilla overview

round

as

and

availability

Western lowland
several sites

at

the Central African Republic (CAR),

in

Gabon

Congo, and

permit.

opportunity

have been studied

gorillas

despite

but,

greater

their

numbers, they are less well known than either

of

the eastern subspecies. They occupy a diverse range

montane

lowland,

Including

habitat types,

of

open or closed

forests with

forests;

swamp, and

arrangement describes almost


and about 60 percent
balance
groups.

made up

is

It

regenerating. The rare Cross River gorilla

gorillas, for

Include

and

stems,

leaves,

fruit,

diet

even

is

known

is

to

Invertebrates,

piths.

generally considered that

Is

It

widely available

in

western

and that

habitat,

gorilla

exceeding 20

lifestyle. Typically

home ranges

fruit

more

is

gorilla than In

this

more

more mobile

km^ western

gorilla

are larger than those of mountain

(except at

Bwlndll,

and there may be

extensive overlap between the ranges of neigh-

boring groups.'

^'"

These overlaps lead

to gorilla

may

inherit or

When
share

to contain related adult

among

usual, however, especially

maturing males

males.

lowland

leave their natal

to

group, either taking females with them, spending

time

In

an all-male group, or remaining solitary


can establish a group

of their

own

by

attracting females.

Females also transfer between groups, some-

eastern

accounts for their

greater fruglvory and the associated

gorillas

is

until they

soil."

their natal groups.

in

mating rights within that group. Hence, multimale

groups are believed

its

mountain groups. The

all-male and multimale

silverbacks they

and forests that have been disturbed and are

although

gorilla

multimale groups result from males

llost

become

they

of

of

maturing and remaining

canopies; forests with dense or sparse understorys;

less well studied,

western

all

groups, about 90 percent of eastern lowland groups,

times more than once.


her

If

a female has an Infant with

at the time, there Is a

being

group.''^
this.

It

Is

serious risk of the infant

by the dominant

killed

male

therefore hard to see

Several factors are

predisposition

to

leave

likely to

the

of

be influential: a

group,

natal

preferences, and her aversions

new

the

why females do

may

all

her

influence a

mate. Following the death

groups sometimes encountering one another. Such

females choice

occasions Involve vocalizations and chestbeats from

adult male, either the females transfer to one or

both groups or only one of the groups, and can lead

more

to

one group moving away. Encounters are some-

may

times violent and

involve lethal

wounding.

of

When

Mountain gorillas

in

taken over

Bwindi National Park,

the dominant male of a

Uganda.

different groups, or the

by another male.
Martha

harem

of the

Is

M Robbms

SOCIETY AND PSYCHOLOGY


Gorillas are considered infants until they are

weaned

about three years," or possibly later

western

at

lowland gorillas."

number

into a

''

In

Young or Immature animals

fall

of categories:^' juveniles 13-6 years],

subadults (6-8 years], and young mature males or


adolescents,
years].

and

commonly known as blackbacks

The process

sagittal crest of

(8-11

on the back

of silvering of hair

mature males (age

12+,

known

as sitverbacks] begins at 10-11 years of age and

is

completed by about 15-16 years. Females do not

undergo

this silvering

maximum

lifespan

become

as they

of

gorillas

in

adult.

the

The

wild

is

unknown,^' with the oldest known mountain gorillas


at over
In

AO years, while the oldest

gorilla to

have died

a zoo reached 53 years of age.'

Median group sizes

of

between seven and 16

animals, most being typically between eight and


11,

have been

gorillas,
diet.

reported for

all

populations of

regardless of habitat type and prevailing

This median represents a dominant, sllverback

adult male, three or four females, and four or five


of their offspring.''

This simple harem-like

99

World Atlas

of Great Apes and their Conservation

muUimale group

dies, the

with the heir, with

whom

social

Gorilla

mothers and

females tend

remain

bonds, except those between

their infants,

particularly strong.

to

they are already familiar.

do not appear

As they get

be

to

older, infants

periphery. Affiliative (friendly or cooperative) inter-

actions between males are therefore rare, but males


in

multimale group occasionally cooperate

The reproduction and sexual behavior

and

juveniles spend an increasing proportion of their

to

prevent females from leaving."

western gorillas

is

little

understood; this

of

one

is

of

they reach

the topics that have been better studied

in

adolescence." Interactions between adult males

gorillas than the other subspecies."'

" " Female

time close

to the

dominant male

and females are largely limited


izations

to

until

exchange

between adults Isee Box

8.3),

displays by males towards females,


of

of vocal-

aggressive

appeasement

males by affected females, and interventions by

males

to

end disputes between females land some-

mountain

gorillas reach sexual maturity at six or

seven years, although between the


estrus-like behavior and the
IS

two year period

amount

moderate aggression and pose

and partner

relationships between the

but

may

to

to

the

males and the females,

limit the effectiveness of

and help males

risk

female coalitions

maintain dominance." Males

multimale groups interact

little

in

with each other, and

of

of

experimentation
situations.

first

bout of

first

conception there

adolescent

chimpanzees and bonobos,

times vice versa]. These interventions involve only


little

mountain

as

sterility;

in

this allows for a certain

among

potential group

The menstrual cycle among

adults has a median length of 28 days, during which

females are most receptive and attractive


ovulation, that

is

one

for

to four

days

around

at

at mid-cycle,

and mating or mating attempts occur

near

at or

relationships between silverbacks and blackbacks

peak estrogen concentrations during menstrual

are generally minimal since the latter are subor-

cycles and pregnancy.^""

dinate and tend to spend a

lot of

time on the group's

single-male mountain gorilla groups, that

In

male

sires all the offspring.' In multimale groups,

Gordon Miller/IRF

males also mate, although often

Gorillas leave a trail of

subordinate

trampled and broken

with less

vegetation behind them.

harassed by dominant males while doing


still

fertile

manage

subadult females."" They are often

to sire a

so,

Mating with individuals from other groups


ceptionally rare.
to

be important

but

proportion of the offspring.^

Female choice

of

among mountain

is

ex-

mate seems

gorillas,

and

is

influenced by male behavior; females either stay


with a mate or leave for another group.

Eastern lowland gorillas share


ductive

characteristics with

including delayed conception, age at


of offspring

(eight

many

mountain

repro-

gorillas,

first

delivery

or nine years), and interbirth

interval (around four years).

Like all other great apes, gorillas construct

nests

which

in

human

to

sleep at night, and can learn to use

sign languages with

novel motor skills taught to


all

some
them

facility,

in

as well as

captivity Unlike

other great apes but the Bornean orangutan,

gorillas have never

tools

in

been observed making or using

the wild. This

that tool use


sufficient

is

is

consistent with the notion

linked to sociability

intelligence

broadens the pool

and learning
of

potentially

in

animals of

ability,

as

it

discoverable

and learnable behaviors. Solitary animals that

seldom meet (such as Bornean orangutans) and


group-living

animals that seldom interact with

one another (such as


likely to

100

gorillas)

should be the least

develop the use of tools. An alternative

Gorilla overview

The history

of

conservation

is

peppered with

conflict,

and park rangers work


hard to maintain the

good

will of their

communities, as here
in

Uganda.

Gordon Miller/IRF

explanation

is

that gorillas are not typically chal-

eastern lowland gorilla during this period, while

lenged by their foods, so have had less need to

armies, rebels, refugees, and miners

develop tool use.

the land. Bushmeat, including that of the gorilla,


still

consumed

in

great quantities.

all lived off

In

is

May-June

CONSERVATION CONCERNS

200^, the rebel military occupation of Bukavu, and

Of the eastern gorillas, the mountain subspecies

the accompanying destruction of equipment at the

has very small but stable populations

Tshivanga

managed
gorillas

national parks

in

iOO

320 gorillas

in

in

l<m' of forestl'"

200 l<m'

in

several well

the Virungas (about 380

and Bwindi (about

of forest)."

The parks are

station

field

showed

Park,

was

not yet stable.

has an estimated total surviving population of

250-280

mental organizations and the

10 highland areas.'^

community,

Kahuzi-Biega National

Of the western gorillas, the Cross River gorilla

well supported by both international nongovernscientific

in

that the situation

fragmented across more than

individuals,

It

is difficult to

assess popu-

and trends among the much more

by profitable gorilla-based tourism programs, and

lation status

by the governments of the region. These popula-

widespread and abundant western lowland

tions are too small to

meet some

theoretical criteria

for genetic health, are vulnerable to catastrophic

areas,

and new areas

events such as outbreaks of disease, and would

been

quickly be reduced by poaching

and Sumatran orangutan,

if

the vigilance of

identified.

conservationists were to be relaxed. Nevertheless,

Conservation

they are being well cared tor

Specialist

The eastern lowland


immediate concern;
to be

around 17 000

its

in

gorilla

is

population

much greater
was estimated

of

the mid-1990s,'' but

it

is

primate

has been

it

and

International
of

lUCN-The

of the world's

V\/orld

large

have recently

Together with the mountain

Group

Union as one

habitat

of

gorilla,

made across

as censuses have not yet been

the

gorilla

listed

by

Primate

Conservation

25 most endangered

taxa.^

Western lowland

gorillas

are widely distri-

feared that thousands had been killed by hunters

buted across a large forested region and occur

by 200A." Warfare engulfed the whole range of the

numerous protected

in

areas,'" but they nevertheless

101

World Atlas

of Great Apes and their Conservation

numbers

Large
remain

in

ern gorillas
forests, a

of

western gorillas may

Congo Basin. The presence

the

substantial densities

living at

west-

of

swamp

in

widespread habitat that was previously

considered unsuitable, was only confirmed


1990s' " after

reports

first

Gabon

national boundaries,

1983.

in

the

in

terms

In

of

thought to hold the

is

largest populations of western gorillas, probably

followed by Congo.

estimated

In

the early 1980s, there were

be 40 000 western lowland gorillas"

to

which about 35 000 were

swamp

discovery that western gorillas also inhabit


forest

numbers, subsequent

significant

in

of

Gabon." After the

in

total

population estimates were revised upwards to

500-1 10

9/4

were made

000.'' '*

^'

However, these estimates

prior to the significant recent impact of

both bushmeat hunting and the Ebola virus.

While additional knowledge

guide

will help to

conservation action, the long-term survival of the

western

gorilla

an increasingly disturbed and

in

human-dominated landscape must depend on the


attitudes of local people and the partnerships they

establish with

government and conservationists.

Modern approaches

to

conservation therefore focus

em-

on community engagement, education, and

powerment, as well as global monitoring;

implemented

all

whose

advanced and promoted

The basic concept

is

and sustainable development

that conservation

initiatives

more successful where communities are


Great ape conservation will suffer
are

weak

either

is

partnership with governments

in

policy priorities are

accordingly.

this

if

be

will

stronger.

communities

or fragmented, and their interests are

overwhelmed by outside factors or ignored

the decision-making processes of people far

in

away

Current conservation projects therefore typically

propose simultaneously starting with communities


Martha

Robbins

to

Mountain gorilla

infants,

face an

uncertain future simply because of the


scale and

cumulative nature

Virunga National Park,

increasing

Democratic Republic of

threats operating upon them. These include forest

the Congo (topi, and a

clearance for farming, forest fragmentation due

silverback male, Bwindi

to

Impenetrable National

degradation

Park, Uganda.

disease. Hunting and disease are increasing as risk

clearance and the

factors because
forest

areas

roads,

forest

hunting for food, and

human access

is

of

the

to

formerly remote

expanding through logging and

outwards' into the rest

community

...

the primary

poorer inhabitants
a social, political

former stronghold

gorilla," as a

this

and the Ebola

virus,

shows how

combination of factors can be.

is

for the

and

of African

opening of friendly

the people,

values,

result of hunting

need

and enthusiasm

in

western

society,

As has been written

and a sharing

population between 1983 and 2000


of the

level.

of

'inwards' towards the

dialogues, partnerships

problems

Gabon, the

work

conservation:

settlement. The estimated halving of the great ape

dangerous

102

building

logging,

by

of

work

with governments to

in

with

of knowledge

especially the

Conservation here

is

and human problem

...

...

communication,

in

primarily
it

poses

education and

because there are huge dislocations

in
in

understanding. Starting from the bottom, links have


to

be

made between

the

various tiers of rural

communities, old and new, national citizenries and

Gorilla overview

an external public that wants

to

help reconcile

solitary males.

WHAT WE DO NOT KNOW

tions

For the western

For the eastern lowland


in

knowledge

gorilla, the principal

relate to the actual

gaps

listically

wild.
is

DRC, and
be taken

From

promote

to

more

known about

mountain
of

to the actions that

their survival

lowland

is

life

the

popu-

from a social and

bonds that

social

maintain the species' societies, the reproductive

the

development

of

and

individuals,

information such as lifespan

annual home range

first birth.

This information

lowland tropical

population

viability.

in

of

a dearth of information on

perspective,

ecological

ecology or demography of

gorillas

There

the detail of western gorilla

scientific perspective, very little

the

gorillas in Bwindi, the

eastern

in

fundamental ques-

gorilla,

and the actual mechanisms

lation decline.

might rea-

of

remain regarding their numbers and

also

distribution,

numbers and

distribution of the species in the war-torn eastern

part of

and the behavior

forest, the lifespan in the wild,

conservation with development"

is

demographic

the wild and age at

in

crucial

in

assessing

Arluke,

A.,

eds 120011

FURTHER READING
Beck, B.B., Stoinski,

T.S.,

Hutchins, M., Maple,

T.L.,

Norton,

B.,

Rowan,

A.,

Stevens,

E.F.,

Great Apes and Humans: The Ethics of Coexistence. Smithsonian Institution Press, Washington, DC.
Harcourt, A.H. 119861 Gorilla conservation: anatomy of a campaign.
Self-sustaining Populations. Springer-Verlag,

Kingdon,

J.

New York.

In:

Benirschke,

K., ed..

Primates: The Road

to

pp. 31-46.

119901 Island Africa: The Evolution of Africa's Rare Animals

and

Plants. Collins, London.

Dates, J.R, McFarland, K.L., Groves, J.L., Bergl, R.A.. Linder, J.M., Disotell, T.R. 120021 The Cross River gorilla:

natural history and status of a neglected and critically endangered subspecies.


M.L., eds. Gorilla Biology:

Muttidisciplinary Perspective.

In;

Taylor, A.B., Goldsmith,

Cambridge University Press, Cambridge, UK. pp

472-A97.
Olejniczak, C. 120011 The 21st century gorilla: progress or perish?

In:

Brookfield Zoo, The Apes: Challenges for

Conference proceedings. Chicago Zoological Society, Brookfield,

the 21st Century.

http://www.

Illinois,

brookfield2oo.org/content0.asp?pagelD=773. pp. 36-42.

Robblns, M.M. 120011 Variation


Sicotte,

P.,

Stewart,

the social system of mountain gorillas: the male perspective.

in

K.J.,

eds,

In:

Robbins, M.M.,

t^ountain Gorillas: Three Decades of Research at Karisol<e. Cambridge

University Press, Cambridge, UK. pp. 29-56.

Robblns, M.M., Sicotte,

P.,

Stewart, K.J., eds 120011 Ivlountain Gorillas: Three Decades of Research at Karisoke.

Cambridge University Press, Cambridge, UK.


Taylor, A.B., Goldsmith, M.L.,

eds 120021

Gorilla Biology:

Muttidisciplinary Perspective.

Cambridge University

Press, Cambridge, UK.


Tutin, C.E.G. 12001) Saving the gorillas [Gorilla g. gorilla]

and chimpanzees IPan

(.

troglodytes] of the

Congo Basin.

Reproduction, Fertility and Development ^2: 469-476.


Vigilant, L., Bradley, B.J. (20041 Genetic variation in gorillas.

American Journal

of Primatology

bit.

161-172.

ACKNOWLEDGMENTS
Many thanks

to Colin

Groves [Australian National Universityl, Alexander Harcourt [University of California, Davis),

Martha M. Robbins IMax Planck


of Stirling],

this

Institute for Evolutionary Anthropology!,

and David Woodruff [University

of California,

San Diego!

Elizabeth A. Williamson [University

for their valuable

comments on

the draft of

chapter

AUTHORS
UNEP World Conservation Monitoring Centre
UNEP World Conservation Monitoring Centre

Julian Caldecott,

Sarah Ferriss,

103

World Atlas

of Great Apes and their Conservation

TJ. Rich/natureplxom

106

Western gorilla [Gorilla

gorilla)

Chapter 7

Western

gorilla

[Gorilla gorilla]
Sarah Ferriss

western

The

18^7)

gorilla

Savage,

BEHAVIOR AND ECOLOGY

broad chest

Observational challenges

iGorilta gorilla

a large animal, with a

Is

and shoulders, a large head, and a hairless,


shiny black

weigh up

to

^-

" Full-grown adult males

about 180 kg, about twice the weight

females. Two

adult

of

face.^^'

subspecies have been

described:" the western lowland


gorilla Savage,
[G. g. diehli

gorilla

[G.

and the Cross River

18'17|

g.

gorilla

Matschie, 190^). The western lowland

much more widespread and numerous

gorilla is

than the Cross River gorilla, which

restricted to a

is

Most research on

gorilla

ecology and behavior has

focused on eastern gorillas [Gorilla beringei],


particularly the
of

mountain

gorillas (G.

the Virunga Volcanoes

Uganda;

been

has

there

comparatively

research on western gorillas

many differences between


many questions remain. By the

beginning of this century, researchers at only three


study sites had succeeded

in

boundary. This chapter will focus on the western

lowland gorillas for study'

^'

lowland gorilla: the Cross River gorilla

ficult

in

Box

discussed

is

7.1.

little

Recent

gorilla].

16.

research has identified


the two species, but

small area on the Nigeria-Cameroon

relatively

beringei]

b.

DRC, Rwanda, and

in

in

because

habituating western
" Habituation is dif-

and

of the limited mobility

the dense forest, the large

home ranges

visibility

of

west-

ern gorillas, and because the gorillas often flee at

DISTRIBUTION

the approach of

Western lowland gorillas are widespread through-

hunting

out

West and Central

Africa

Congo/Oubangui River seems


boundary

of

boundary

is

the

(see

Map

their distribution,

essentially defined by the course of

the southern edge of their distribution


by the forest-savanna boundary, as
7.1.

Western

their

of

formed by the Atlantic coast, and

is

gorillas

Cabinda province

of

are found

humans due

the

past.

to having

experienced

However, some excellent

observations of less habituated gorillas have been

made

in

where

marshy forest clearings

visibility is

typical of that

good. Not

seen

in

alt

(locally called bais],

behavior

bais

in

is

forested environments."

limits of the

The western boundary

forest.

distribution

eastern

and the northern

Sanaga River and the northern

closed

The

7.11.

to delimit the

in

defined

is

shown on Map
in

Gabon, the

Angola, the western part

Congo, the extreme southwestern part

of

of

the

Habitat

Western lowland
growth)

and

(including

gorillas

occur

in

primary

secondary (regenerating)

forest

swamps) as well as

submontane" and lowland

lold

forests
in

both

areas.' Overall, western

lowland gorilla occurrence, biomass, and density

seem

to

be positively correlated with terrestrial

herbaceous vegetation, particularly

Central African Republic ICARl, south-central and

or aquatic

southern Cameroon, and

monocotyledonous plants [including gingers and

Guinea.

They used

western

tip

of

the

to

in

mainland Equatorial

occur

in

Congo IDRCl, but are now probably


country.

the

extreme

Democratic Republic

of

the

extinct in that

palms)."-'"'"'

During the 1980s, Tutin and Fernandez found

western gorillas
surveyed

in

in

seven

of

15

habitat

types

Gabon:''" dense primary forest; dense

105

World Atlas

Map

7.1

of Great Apes and their Conservation

Western lowland

""

Box

gorilla distribution (see

'^oE

~ "Ty

7.1 for

Cross River

Data sources are provided at the end of

gorilla)

IS-E

--

'-'

J^,

Species

MOUNTAINS

IrO

l^-ZJ",

Western lowland

gorilla

Unidentified gonlla species

Crass

gorilla

Estimated range
Western lowland

"RiverNP

chapter

Confirmed range

Western lowland

CAMEROON

this

gorilla

^y

-^

v^

CENTRAL
AFRICAN REPUBLIC

Polnte-Noire
5'S

5S

CABINDA _

ANGOLA
lO'E

106

bas-fleuve

Western gorilla [Gorilla

inundated (swamp) forest; thicket; undisturbed

secondary vegetation; exploited forest (one


years after logging); exploited forest (two

two

to

years

to six

after logging);

and coastal scrub. Western gorillas

appeared

be absent from areas of

to

human

Although

was

it

now considered important

swamp

in

season." The soils

Reserve (now part

of

In

The

fairly

forests both
of

in

the wet and the dry

these

tend to be

herbs found here, such as Hydrocharis spp.

Parl<),

is

(Hydrocharitaceae), can provide important nutrient

sources for western

swamp

common

Species

gorillas."^

high density of western gorillas seen

genera Xylopia (Annonaceae), Raphia (Arecaceae),

in

the

the Dzanga-Ndol<i National Park

of

attributed to the presence of

which

forest,

is

such as herbs."

rich in nutritious folivore food

Western lowland gorillas have been observed


occasionally nesting along savanna-forest edges or

savanna

itself."-

'"

"^-

Western lowland

forest include those belonging to the

Klaineanthus (Euphorbiaceae),

Lophira (Ochnaceae),

Aframomum

Caesalpinioideae),

and

Zingiberaceae).'"

northern Congo, a study found

that

In

western gorillas favored those

where Raphia was common,


western CAR, the distribution

of forest

fragments within the savanna. They do not

seems

these

in

habitats,

however,

to

be

Aframomum

influenced

(gingers,

swamp

forests

palm genus used

both for food and nest construction."

sometimes,'" but not always,'" make use

permanently

Trichiiia (Meliaceae),

Guibourtia iLeguminosae-

gorillas

live

swamps

where

Loango National

moderately disturbed or secondary

the

high

to

CAR have been

in

in

secondary forest with more edible herbs.'

Dzanga Sector
in

feeding

waterlogged or permanently flooded; the aquatic

generally scarce, western gorilla nests were found


in

habitats and

the Petit Loango

the herbaceous vegetation favored by gorillas

mainly

that

forests are

densities.""^ Western gorillas have been observed

avoided roads and plantations, but were observed


recovering secondary forests.

thought

swamp

areas for western gorillas, supporting them

settlement and disturbed secondary forests; they


in

originally

western gorillas avoided water,

gorilla]

of

In

south-

western gorillas

by the

availability

of

spp.^'

perhaps because the forests do not provide either

A mate western lowland

sufficient preferred nesting materials or a constant

Diet

food supply'"

There are two major differences

The Odzala-Koukoua National Park

in

Congo

between the habitat

of the

provides a good example of the variety of habitats

of the best-studied gorilla

occupied by western lowland gorillas. Here, they

Richard Parnell

primarily
In

which

live

in

open-canopy Marantaceae

sufficient light

food availability

gorilla feeding,

and that

Nouabale-Ndoki

subspecies, the mount-

National Park, Congo.

in

western

gorilla

forest.

reaches the forest floor

to

allow plentiful understory vegetation to develop."


This forest type

dominant

is

to the north of the Lekoli

River, particularly in the northeastern part of the

park.

The ground vegetation

most impenetrable
including

is

thicket of

Haumania

dominated by an

al-

Marantaceae species,

Megaphrynium

liebrechtsiana,

macrostachyum, and Sarcophrynium spp. Western


gorillas are also found in the

closed-canopy primary

forests of the park, which have a

much more

con-

tinuous canopy and a sparser understory.


In

and around the northern part

of the Odzala-

Koukoua National Park there are more than 100


forest clearings.

Those that have been investigated

have a particularly sodium-rich marshy herbaceous


vegetation and are

Box

7.2).

Western

known as

salines'" or bais (see

gorillas are

known

to visit

these

clearings daily to feed on plants from families such

as Cyperaceae and Asteraceae, which here are rich


in

sodium absorbed from the

soil." Bais therefore

provide a unique opportunity for researchers to

observe the animals

in

the open.

107

World Atlas

of Great Apes and their Conservation

ain gorilla, affecting botfi diet

and foraging behavior.

First, 'tiigli-quality' tierbs that

and

rich in proteins

are easily digestible

and minerals are much less

abundant and more patchily distributed

swampy

gorilla habitat, outside


is

much more widely available

in

in

western

lar

gorilla

available;

group include fecal analysis, and the

show signs

monitoring of food types that

having

of

been processed by western gorillas and are


along gorilla

Western

trails.^'

large

amounts

fruit,

piths, invertebrates,

gorillas

left

consume

and

soil.

There are sea-

sonal, annual, and spatial variations in the frequency


of

consumption

Table

of

items'"' '"

food

different

Isee

The seasonal importance

and herbs

of fruit

in

the diet of the western gorilla has been the subject of

debate."

appears
of

to

January-March, when few fleshy

more

Favored tree

fruit is

fruits

eaten at other times.''

fruits include

" '"^ The

availability of

seasonal

fruit

shape the foraging and ranging patterns

western

seasonally,

those of the genera Tetra-

[Gambeya] [Sapotaceael, Diatium (LeguminosaeCaesalpinioideael, and Landoiphia lApocynaceaeJ.^''"^

The

herbs such as species

fruits of terrestrial

Aframomum, Nauclea

Country

site

gorillas.'"''
it

may

When

constitute

fruit

most

is

phrynium (Marantaceael are also eaten when


available."'"

Some

habitats of the western lowland gorilla

are dominated by the leguminous tree Gilbertio-

dendron dewevrei (Leguminosae-Caesalpinioideael;


roughly

at

five

year intervals this tree produces

High-

The western

of nitrogen.

gorillas feed heavily on the

seeds during these mast


willing

to travel

stands of

abundant

of the diet.

Species

6.

some

fruiting events,

and are

distance to congregate

Fruit

animall

CAR and

incl.

High-quality herbs,

where

available, are im-

portant for western gorillas. Plants from the

mono-

Invertebrate

Other

Leaf

Stem/pith

Flowers

Bark

species

foods'

seeds

100

70

33

yes

138

77

8^

Congo

CAR

Bai Hokou'"^

dirt

from termite

mounds and

bais''

Congo

152

133

29

10

6 (includes roots

Belinga'^'

Gabon

89

72

18

A Inonplant foodsl

Lope National

Gabon

17

8 (includes roots,

16

included

10

22 (includes bark,

Nouabale-Ndoki
National Park"'

and shootsi

Park"'"'
Lope National
Park'"

Gabon

not

91 +

seeds

known

of 21 species

not

100 + seeds

known

of 21

up

to

49

galls,

A8

species

in 'other'

roots,

and

Cameroon

>22

Cameroon"'
Afi

Mountain

Nigeria-

168

Cameroon

Wildlife

Sanctuary'
a Wood, shoots, buds tubers, rhizomes, and fungi,
b See Box

7.2.

c Cross River gorillas


d

108

No evidence

found n fecal samples, feeding

trails

etc

not

not

not

not

not

known

known

known

known

known

TOO

36

22

53

22

0"

not

fungi]

wood,

and
Southeastern

in

dewevrei."

Plant species

Iplant or

Mondika"

of

and Mega-

(Rubiaceael,

Table 7.1 Western gorilla diet

Study

are

"*''''

especially nutritious seeds that contain high levels

7.11.

much

of

pleura ILeguminosae-Mimosoideae), Chrysophyllum

and they eat leaves, stems,

of fiber,

minerals and proteins

all

and woody vegetation are consumed during the dry


season

to identify the diet of a particu-

rich in

year round, while low-quality herbs are


eaten only when fruit is scarce.^' ""^ '"' More leaves

the habitats of west-

the absence of direct observations of feeding,

In

methods used

herbs that are

areas. Second, fruit

ern gorillas, so they eat significantly more fruit than


'"^ '"
do their eastern counterparts.'^' " '"' '"' '"

the

quality

are eaten

soil,

funqil

known

3 (rootsi

'

Western gorilla [Gorilla

Box

7.1

THE CROSS RIVER GORILLA

[GoriUa

mented highland areas." Despite the


dense human population

The

gorillas inhabiting the

that straddles the

Cameroon

at

mountainous landscape

border between Nigeria and

the headwaters of the Cross River

were recently recognized as the subspecies


gorilla diehli.

estimated

more than

individuals distributed across

gorilla diehli]

These Cross River


surviving

total

Gorilla

gorillas have an

population of 250-280

this

10 fragrelatively

region of West

these gorillas have persisted, protected by

Africa,

and

adaptability

their

in

gorilla]

human-development

relative

activities

region and gorilla habitat


survival

future

of

this

is

inaccessibility

As

increase within the


further eroded, the

ape depends on urgent

conservation action.
continued overleaf

Cross River gorilla distribution


9ffE

l^Jjt I A

>-:..N

ktt).*-''^^

(Okwangwo

Divisiori):*

v:4'

cotyledonous Marantaceae family, for example, can


provide a dependable supply of food

all

year round.

Marantaceae genera that are frequently eaten


include

Megaphrynium and Haumania.

If

available

within a group's range, western lowland gorillas

often

feed

on aquatic and semiaquatic sedges

(Cyperaceael and
cordifolia,

include species of Hydrocharis and the sedges

Rhynchospora
In

the

(all

Fimbhstyiis [boih Cyperaceael."

forests of the Likouala region of

northern Congo, the fronds of Raphia palm are

consumed,

along

with

species

of

Pandanus

IPandanaceael and AframomumJ'


Outside fruiting seasons, more fibrous vege-

herbs such as Marantochloa

M. purpurea, and Halopegia azurea

ar\6

swamp

tative

matter

is

eaten, including shoots, young leaves,

the absence

preferred

foods,

Marantaceae], visiting streams, bais, and riverine

and

swamps

Nouabale-

western gorillas eat leaves, bark, low-quality herbs

Ndoki National Parl< ICongol, preferred food plants

such as Palisota ICommelmaceael and Aframomum,

to

do

so.'

'

At Mbeli Bai

in

bark."^

in

of

109

World Atlas

of Great Apes and their Conservation

Cross River gorillas

Geographical distribution

of

Cross River gorillas are the most northern and

Nigerian

western

that,

the

Separated from

of all gorilla populations.

nearest gorilla

approximately 200

population

the south

to

km, they occur

Mountains Community Forest, the

the Cross River National Park,


the

Takamanda

Mountain

Afi

and the Okwangwo

Wildlife Sanctuary,

all in

by

Mbe

the

in

Division of

Nigeria;

and

Forest Reserve (contiguous with

Okwangwol, the Mone Forest Reserve, and the


Mbulu

Community

Hills

Cameroon"

Forest, all

southwest

in

by the late 1970s,

were probably

were described

1904, the gorillas of this region

as a new species: Gorilla diehli Matschie, 1904

in

this led to

Nigeria"' and

in

of gorilla

systematics

amen-

subspecies Gorilla

to that of the

of

1968, Critchley

In

in

1989, Harcourt and

In

coworkers estimated that a further 100-300 gorillas

remained

1996, the

Nigeria, In

in

1997 by a study

in late

extended suivey
of

in

long-term

first

Nigeria,

in

Cameroon. These

the subspecies Gorilla gorilla gorilla Coolidge,


the last years of the 20th century, cranio-

efforts.

The most recent estimate

the Cross River gorilla population, published

2003,

that there are

IS

150

in

Cameroon." Surveys

in

in

between 205 and 250 weaned

70-90 individuals

individuals, with

suggest that

other western gorillas

all

meant

that gorillas

renewed surveys both

Cameroon."

Takamanda, Cameroon.''^"

were amalgamated with

In

was assumed

had estimated that 25-50 gorillas remained

about

1929.

it

extinct in Nigeria,^" During the 1980s,

there;*"

gorillas

gorilla diehli Rothschild, 1904, 1908. In 1929, they

in

The 1966-1970

studies are ongoing, and have been accompanied by

Taxononnic history and status

ded their status

122

77,

study on Cross River gorillas began

(see map],

Subsequent revisions

32,

3.

2,

lack of information

however, reports appeared on the persistence

followed

In

war and

civil

Nigeria and

progress

in

few previously unknown sub-

populations remain to be confirmed, producing a


tentative total population estimate of 250-280.

metric IskuU measurement] studies found that

Cross River

lor 'Nigerian') gorillas differed signifi'"- '"'

cantly from other western gorillas;'"-

their

subspecies status was then reinstated by Groves


primate taxonomy^'

his review of

Despite

new conservation

and the animals are

lost,

at a low level. Given their

in

they

survival,

still

Endangered

in

have
the

listed

List of

as

above sea

is still

area for very

to

their

Critically

lUCN-The World

lowland

inhabit

gorillas

Although people have

small and fragmented

been

Red

River

recent

being hunted

and the continuing threats

population

Cross

semi-

deciduous and evergreen submontane forests from


about 200

efforts

years, the habitat of the Cross River gorilla

being

in

Ecology

many

level to at least

lived in

and around

500 m.

this forest

generations, there remain large

tracts of primary forest throughout the habitat of

the

Cross

River

particularly

gorilla,

Cameroon. Most subpopulations

above 400 m; these are

gorilla exist in ridge forests

more

typically

difficult for

within

Cross River

of

hunters

to

access due

to

the steep terrain. At the highest altitudes across

Conservation Union.'"

the Cross River gorilla range, farming, burning, and

Population

From the

cattle grazing

early 1930s to the late 1960s there

scattered reports on the distribution and

and less-favored

fruit such as Duboscia


and Klainedoxa ISimaroubaceael.' "

Western lowland
seen

to eat at least

one study

in

Most

of the insects

in

Cubitermes sulcifrons

110

in

of

weaver

of

are also

In

Gabon, insect
of feces.'''

the termite

ant (including the

It

in

Cameroon, the

gorilla

more

rarely,

grubs, and larvae from dead wood,

known

is

and

of forest

the edge of the

ant, Oecopliylla longinoda] and,

caterpillars,

30.5 percent of western

Three species

especially on

ITiliaceael

eaten are termites and ants;''

one study found the remnants

gorilla feces.

in

about one third

have produced a mosaic

here,

Bamenda Highlands

have also been

gorillas

grassland;

abundance

20 species of invertebrate.'''

Lope National Park

remains were found

were

to

be eaten."''

"'

possible that western gorillas have a food

culture, with learned preferences

passed on be-

tween individuals and generations.'^ For example,


the

species

culturally.

of

insect

Weaver ants

eaten
(0.

large quantities at a study site


in

Gabon, but

in

appears

to

vary

longinoda] are taken


in

in

Lope National Park

the Belinga study site 250

km

away.

Western gorilla [Gorilla

Kelley McFarland

KeUey McFarland

The Cross River

extremely

gorilla is

catch on camera.

difficult to

habitat consists of relatively small isolated patches

sometimes tenuously connected by

of forest,

gal-

field

Mountain Wildlife Sanctuary," suggests

when

fruit

habitat

is

at

Afi

that, like

five

months.

is

scarce and the


pith of terrestrial

herbs, bark, and leaves.

Cross River gorillas are rare and shy as a

most information on

result of hunting, so

ecology and behavior has

their

come from an exami-

nation of nest sites, feces, and feeding trails. Nest

clusters suggest that group size

(fewer than

occasional

six

much

termite species

weaned

sutcifrons]

[C.

both sites."' Similarly,

eaten by

some

is

typically

individuals],

if

potential

Cross River gorillas are

for

gorilla

Large areas

between many

connectivity

of the

areas need

to

some

despite being available

in

not

exat

species are
by others,

the habitat of both.

to

be strengthened. Along with other conorganizations, the Wildlife Consen/ation

Society

working on both sides

IS

between

diet

is

strong

of the

Nigerian-

in

collaboration with state and

national government

agencies, to improve con-

Cameroon

border,

servation of the gorillas.

Jacqueline

L.

Sunderland-Groves, John

relationship

and foraging behavior Species or

populations feeding on high-energy foods that vary

F.

Dates,

and Richard Bergl

spatially

and seasonally tend

to

have greater day

ranges (average distance traveled by a group per


day! than those feeding on lower-quality but

more

consistently available foods. Western gorillas


this pattern, as they travel farther

shorter day ranges


there

protected

servation

needs

and termites are available

Day ranges

Among primates

new

be created and local law enforcement

small

was eaten

plant

unoccupied

subpopulations.

To maintain these forest corridors,

although

weaver ants occur

populations and

of

remain; these provide

habitat

larger groups do occur Nesting

clusively instead, although

some-

they and their habitat can be ade-

quately protected.

dominated by the

is

will

Conservation prospects

But the Cross River

fruit

about 20 individuals

The survival prospects

available.

notable for strong seasonality,

lAugust-Januaryl,

gorilla diet

of

smaller foraging parties."

promising

the late wet season through early dry season

at Afi

into

diehli preferentially

6.

with a dry season that lasts four to

From

group

g.

other vtfestern gorillas,

gorilla

Afi

study yet completed

on Cross River gorillas, by McFarland

feeds on

Mountain Wildlife Sanctuary suggest

patterns at
that a large

times divide

lery forest.

The only long-term

gorilla]

and woody

when

vegetation.^'

much

must
fit

rely

of

on leaves

this pattern in

gorillas, as their

greater than that

fit

fruit

the forest, and have

They also

comparison with eastern


is

in

they

when more

either

day range

mountain

111

World Atlas

of Great Apes and their Conservation

year period found that western gorilla ranges varied


in

between years, with the estimated minimum

size

home range

A western

being 22.9 kml'"'

gorilla

group observed at Lope National Park, Gabon,

km' over 10 years, but probably

visited 21.7

cover the entire area each

did not

The home ranges

western gorillas are less affected by

of habituated

human

year''"'

presence, so caution

comparing data from

and position

shifts in both the size

range also occur, probably


seasonal

availability of

much smaller

response

in

fruit,'''

Temporary

home

of the

the

to

and ranges may be

during the dry season than during

when

the wet season,


but nutritious

needed when

is

different studies.'^

the gorillas feed on dispersed

fruits.'"'

Ecological role

Western

animals that

gorillas are large, dexterous

affect the structure

and composition

of vegetation

by feeding on plants and building nests with them.

Trees can be badly damaged by gorillas climbing

and feeding
Iroko Foundation

The buttress

of a large

tree iParkia bicolor)

growing

in

the Oban

gorillas

amid abundant lierbaceous foods or

eastern lowland gorillas


fruit availability;

there

is

living in

of

places with poor

no such difference between

Division of Cross River

the day ranges of eastern and western gorillas

National Park.

when

both

live in

areas rich

'^

in fruit."'

At Lope National Park, Gabon, the western

lowland gorillas travel about


Bai

Hokou

between

about
2

with an overall

km/day

mean

sufficient food.'"'

it,

day.'^' At

in

'"'

order

Larger

to obtain

presence, and the degree


to

human

hunters and leopards [Panthera pardus] have been


to

influence the
gorilla

the

vicinity,

movement

patterns of

when predators

presumably as a result

avoid detection or to evade

fruits with

them

are

of efforts to

seeds

intact,

seed dispersers, with up

samples containing seeds; samples

cally contain

many

Seeds deposited

seeds, most of

at

99 percent of

to

them

typi-

intact.

appear

some

to

be particularly important dispersers

species, for example the

endemic Cola

area used by a group

[Sterculiaceael of Gabon, the fruits of which are

consumed
after

avidly

and have a high germination rate

deposition.'

means, western

Over time, by these various

gorillas might have a significant

influence on the forest.

A number
same
its

of

other species use

some

of the

food resources as the western gorilla and share

habitat, so competition

between them may occur

is

particularly

gorillas

interesting;

CAR,

Congo,

Equatorial

Guinea,

aggressive interactions between

those of mountain and eastern lowland

been observed."

gorillas;'"'

home ranges of different groups overlap


extensively.'" A study at Bai Hokou, CAR, over

quite
a

two

of

tizae

and

although

they have overlapping diets and are sympatric


(the

'^

nest sites have higher ger-

over a year) of western gorillas are larger than

the

leaving only

all fecal

The relationship between western

entirely."

Home ranges

112

their

gorillas are

and swallow

effective

chimpanzees

The annual home ranges

" Western

certain trees, '^'

to

those seeds that are too hard or too large. "^ They are

groups. Very long or very

short distances are traveled


in

most

visitors

mination and survival rates than those deposited

also affects daily travel distance.'^ Both

some western

reliable

elsewhere, such as on paths."" Western gorillas

which the western gorillas are accustomed

observed

increasing that of others."

the

plant species, and

frugivorous

of 2.3-2.6 km/day*"'

Human

some

reproductive success of

folivorous months,

during

km/day during

groups travel greater distances

to

each

CAR, the distance traveled varies

in

months, and

km

them.'" Like other frugivores, they

in

consume and disperse seeds, reducing

both

When

this issue

Belinga area of Gabon, and


National Park

in

in

and

in

Gabon,'"

them have never

was

studied

in

the

the Nouabale-Ndoki

Congo, 60 percent and 42 percent

Western gorilla [Gorilla

Box 7.2 FOREST CLEARINGS: A WINDOW INTO


THE WORLD OF GORILLAS

considerably from

Residents
It

to

IS difficult

observe western lowland gorillas


as

their rain-forest fiabitat,

through

little light filters

the canopy, and the understory

dense vegetation. Meeting

is

in

often choked with

gorillas

circumstances can be alarming

under such

humans and

for

gorillas alike; contacts often Induce aggression or


flight,

so collecting unbiased behavioral data

almost impossible.

knowledge

based almost

Then
clearings
bai5:

in

Yet.

western

of

is

until the early 1990s,

our

behavior

was

gorilla social

the forest, which local people called

these appeared

gorillas into the open,

to

western lowland

attract

and offered

a fresh

their world. Several studies of bais

quently undertaken

understanding

of

view

of

were subse-

Congo, CAR, and Gabon.

in

Together these have contributed greatly


the

lives

of

our

to

western lowland

The

classic gorilla bai

is

from 0.04 km^

llboundji,

Congo]

a treeless clearing

The

size

may

vary

about 0.18 km^

to

IMaya Nord, Congo!; most are either roughly


circular or linear

in

form. The substrate

is

often

extremely swampy, with water-saturated soils


loosely held together by a

Small streams and


although the area

An openly exposed

mat

pools

of floating vegetation.

are

often

of relatively dry

present,

many

other species.

Congo swamp

otters \Aonyx

antelopes [Tragelaphus spekei],

sitatunga

eagles iHaliaeetus
umbretta], and
Sa/visitors

lily

may

vocifer],

fish

hammerkops IScopus

trotters lActophilornis afr/canusl.

include forest elephants [Loxodonta

cyclotis], forest buffalo

river

[Syncerus caffer nanus], red

hogs iPotamochoerus porcus], giant forest

hogs [Hylochoerus meinertzhagenil. and guereza


colobus monkeys IColobus guereza].
activity of

western gorillas while

at bais is feeding. This generally

begins as soon as

an individual or group enters the


shortly before they leave.

It

is

primary reason for western gorillas

to

in

protein,

to

is

usually

Cyperaceae, and Gramineae

bai plants are highly digestible, high

and contain elevated

other minerals (attributes that


tive

western

gorillas).

levels of salt

make them

bai plants

is likely to

ingested

at

Despite these qualities,

limit the quantity that

single

and

attrac-

the high water content and low dry weight of

the

is

dominated by plants from the

Hydrocharitaceae,

Many

imagine,

to visit bais.

Aquatic and semiaquatic vegetation

abundant, and

and ends

bai,

easy

availability or quality

therefore, that food

families.

gorillas.

situated around a watercourse.

include

The primary

marshy

discovered

may

Western lowland

bai.

to

congica], spot-necked otters [Lutra maculicollis],

on such observations.

entirely

primatologists

bai

gorillas share these areas with

gorilla]

sitting.

At

Mbell

many

can be
Bai

in

Nouabale-Ndoki National Park ICongol. the average length

of visits by

groups was only two hours.

ground varies

continued overleaf

family group of western lowland gorillas and a silverback male.


Richard ParnelL

Richard Parnell

113

World Atlas

of Great Apes and their Conservation

Much beyond
too

and the

this,

When some

species of bai plant Isuch as


chevalieri,

the

of

Hydrocharitaceael are picked, they are often

lightly

coated with sediment. Western lowland gorillas

wash

sediment fastidiously by waving each

off this

handful of plants back and forth


eating

western gorillas

wade
1

in

the water before

To reach this succulent food source,

it.

walk through swamps and

will

bipedally across streams that are

deep,

using

outstretched

their

balance. Until quite recently,

made

gorillas

point

it

western

gorilla

may

more than two hours

for

seem

that

at a time.

home

at

feeding at a

water up

sit in

for

contact with

When

semiaquatic environment.

bai, a

arms

to their

Even

stream crossings,

will happily

is

dominance

groups or leave

both

paddle and play

in

but. in
tool'

western gorillas may leap bodily

use

splash

bow

slap the water's surface with one or

hands. At Mbeli

Bai,

all

ages

of

western

females, have been seen to

displays;

prompted the

has

this

hierarchy

tolerance

western gorillas

to bais

be any

to

'ownership' of a specific bai by a particular group


of gorillas:

groups or solitary males with

ranges close
visitors,

to a bai are likely to

be more frequent

is

shown

different

groups

by groups toward each other At

led to

actual physical contact


display behavior

groups were

several times a

particular bai

week, while others

may pass

through that bai only twice a year


Bais have

finally

relaxed
forest.

life

that

would

otherwise probably have remained undocumented


for

much

groups
Bai,

of

longer'- Bais can attract

western

gorillas.

many

different

For instance, at Iboundji

researchers have identified i7 different groups

and 25

solitary males.

Sample

only

mouncontact

in

six

times less

to

likely

gorilla

when

engage
It

sizes such as these

in

appears

groups are generally more

close together

The unparalleled

bais than

in

the

in

offered by the bai

visibility

habitat permits a silverback to survey his group's

females closely while monitoring the behavior

of

nearby silverbacks. The resulting reduction

in

that

may

members to

be unthinkable

males practice displaying

in

in

interact in

the forest. Young

front of

females from

other groups without risk of attack, and juveniles

from

groups wrestle and play with each

different

other,

them

potentially
into adult

making contacts

life,

and shape

that will last

their interactions in

the future.
of

western gorillas

we

still

in

bais

have a great deal

themselves, however,

may

is in its

early

to learn.

Bais

provoke or permit beha-

viors that are rare or absent in the

more usual forest

habitat of the western gorilla. At Mbeli Bai,

permitted primatologists to

study aspects of western gorilla

58

7 percent of group encounters; their

western

that

Study

contrast,

peaceful mingling than those at Mbeli.'"

stages, and

visit

by

this

In

was never witnessed;

was observed. By

have been seen to engage

in

home

Some groups may

of bais

an aggressive response;

almost always came from the silverbacks.

therefore likely to be a function of the

proximity of a bai to the center of a group's

range.

home

but no group has exclusive access. Visit

frequency

examine intergroup

Mbeli Bai. only 30 percent of shared uses

ways

to intimidate rivals."

There does not appear

their

of

and how a silverback

to

males

vary widely

females,

behavior One surprise has been the degree of

tension permits other group

visitation rates of

between

of females.

Bais also allow us

suggestion that the displays are used primarily by

The

able to study social

other completely. Where aggression occurred.

streams, generating spray and a large

gorillas, except adult

influences the fate of

now

infants,

places, use water as a communicative

may

are

percent of cases, groups appeared to ignore each

dramatic 'splash displays'. During aggressive or

wave, or

size

voluntarily,

harem

retains his

aggression

into

We

whether young males are pushed out

tain gorillas

playful displays,

the

of

including

life,

dynamics within a group, exploring whether there

shallow water while their mothers feed. Not only do

some

1U

how group

individual gorillas.

western gorillas tolerate water while feeding

via

gorilla social

chest

to its

mothers' backs during

although clinging

western

of

the formation, evolution, and fission of groups, as

well as

more than

was thought

avoiding

of

water, but western lowland gorillas


in

have permitted a more thorough study

dynamics

Hydrochans

aquatic

the

may simply be

gorillas

to continue.

'full'

it

is

estimated that on average, western gorillas spend


only

percent of their daylight hours

in

the clearing.

Bais offer us perhaps the best opportunity


to

we

view the intricacies of western gorilla social

have
life;

however we must also be cautious

in

from what

other habitats.

is

learned to behavior

in

extrapolating

Richard Parneli

Western gorilla {Gorilla

respectively of

all

foods recorded for western gorillas

were also eaten by chimpanzees."'


shared foods are seasonal
plentiful at the

time

of

fruits,

'"

Many

which are

occurrence.

is

It

of the

also

is

differentiation

limited

and by the

when

by each species
(see Box

study sites."'

Western

lowland

which

its

silverback

a shortage of

is

food'''^

occasion.'

few natural predators,

However, western gorillas do not

between

'

sions been suspected of having killed gorillas;"

been
'fear

observed

odor'

scared gorillas! has been detected

(the

seems

predator

therefore,

likely,

that

fleeing

scent

of

leopards.*^'

'"

western gorillas

leopards as dangerous, and that this

may

represent a real threat

to

out with distances of over 500

there

is

more than one

gorillas

almost always contain only one dominant,

gorillas

is

habitats occupied

silver-

back adult, plus three or four females, and four or


five

offspring."

""

Groups that contain more than

one silverback have only very occasionally been


ported
Ian

among western

lowland gorillas,^^""'

re-

'^'

in

Where

subgroups

'^

lowland

similar not only to that of both eastern

is

subspecies, but

size

reproductively active western

"''

among western

also similar across the range of


52, 92, 108, 150
'^

The very large groups

sometimes observed among eastern

them.

Social behavior
of

overnight.^'"'

general, group size

In

up during

split

silverback, these

sometimes remain apart

ever, rarely occur in

Groups

groups

individuals; other

the day and then reunite at the nest site."

the air after

in

encounters between western gorillas and


It

ousted by another from

be as cohesive on a daily basis as


their eastern counterparts.^'- " In some groups,

members spread

perceive

group

of a

to

large and strong. Leopards have on several occa-

leopards," and gorilla

is

form

generally

The takeover

group fission has been reported on only one

gorillas have

have

'''

different strategies applied

there

appear

gorillas

only

outside the group has never been reported, and

as their juveniles are well protected and adults are

western

gorillas

stable cohesive groups."


in

at

^"''''

by other forms of niche

8.11.

Western

some

relatively

possible that

competition between western gorillas and chimp-

anzees

and all-male groups have been reported

gorilla]

western

gorillas,

gorillas."

western gorillas appears

to

'''

'"

how-

Group

be influenced by

the size of the foraging patch, fruit abundance,'"'

and the degree


within

the

siderably

of

competition for food that occurs

group.'" Western

more

fruit

gorillas

than eastern

eat con-

gorillas;

this

dependence on clumped food resources may


constrain their group size. Pressure from predators

Redmond/UNESCO

Young gorillas interact


through play.

115

World Atlas

of Great Apes and their Conservation

Box 7.3 POTENTIAL MEDICINAL VALUE OF


GORILLA FOODS

discovered iboga.

A growing body

suggests that primates

describes boars, porcupines, and western gorillas

and

digging up and eating the roots, and afterwards go-

use certain plants

for tfie control of illness

chimp-

the regulation of reproduction."

In Africa,

anzees and humans show strong

similarities

the

in

Few

plants they use to treat similar symptoms."

equivalent data are yet available for western or

eastern gorillas.

An ethnographic and pharmaco-

logical study of the properties of plants ingested by

and eastern gorillas evaluated their

western

potential medicinal value."

was compiled

list

of

ing into a wild frenzy,

known both
African

ditional

be used by

to

medicine" and

western and eastern gorillas

included antiparasitic,

tra-

in

be eaten

to

by

the wild." Major

in

antifungal,

chimpanzees

number

of

and seeds

of

Cola species ISterculiaceaeJ, notably

pachycarpa which

of C.

Gabon as

gorillas ingest the fruit

people

by

known

is

locally in

The same seeds are

'cola of the gorillas'.

widely chewed

West and Central

in

example by long-distance truck drivers

Africa, for

the

Both

alchornine and

alkaloids

alchornidine,"- and local people use the pith and

leaves as an antiseptic and cough remedy.'^^

The Virunga mountain


visit

gorillas

periodically

upper mountain slopes characterized by giant

Senecio (Compositael plants.

George

1963,

In

Schaller followed a group to an altitude of 4 100

preferring

to

important

in

eat the

Senecio species are

pith.

ethnomedicine

in

the treatment of
colds.

Gorillas

also occasionally feed upon the giant Lobelia plants


that

grow

members

these

at

giberroa and

L.

such as

altitudes,

genus contain

this

of

high

L.

woUastonii ICampanulaceael. All


bitter-tasting

alkaloids that have stimulating effects lasting up to


a

quarter of an

hour One

these alkaloids,

of

in

order to stay awake, and are used commercially

lobeline,

in

cola drinks. They contain caffeine (2-2.5 percent

willingness of these eastern gorillas to travel to

dry weight!

and theobromine, but only small

amounts

protein,

of

gorillas eat

suggesting that western

them mainly

for the stimulating effect

In

Gabon, western gorillas have been reported

to eat the fruit,

stem, and root

Tabernanthe iboga

of

(Apocynaceae). The iboga shrub

documented hallucinogenic
active principle

in

T.

iboga

plant

in

Africa.'"'

The

ibogaine, with the

is

the roots.

in

Ibogaine affects the central nervous system as a

as well has having a stamina-

hallucinogenic,
effect

on the cardiovascular system

similar to that induced by caffeine.


integral

cults

and

component
rites.

region are

also

respiratory stimulant.

The

higher altitudes, expending considerable energy

in

the process, to reach a zone with fewer foods, less

oxygen, and colder temperatures, suggests that the

in

certain

The inhabitants

among

those

to

of

use

once reaching

T.

iboga

African

Is

an

religious

the Petit Loango


T.

iboga for this

purpose. According to Bwiti Ithe religion practiced by


the (vlitsogos and the Fangs, two ethnic groups

this destination are of

consume
some special

value to them.

perhaps the best

is

highest concentration being found

boosting

is

pharmacologically active plants that they

of caffeine.

More

detailed field studies are required to

allow a critical evaluation of the possibility that


gorillas are self-medicating.

significance of the

and

consumption

of

to

explore the

these plants

In

their daily lives. Observations already recorded pro-

vide an important starting point, paving the


a greater

understanding

adaptations of gorillas
This field of research
quality

new

of the dietary
in

may

and population

way

to

needs and

their various

habitats.

help us to assess habitat

viability,

as well as providing

insights into the control or cure of diseases

that afflict both

apes and humans.

in

Gabon! legend, the forest-dwelling pygmy' peoples

116

cordifolia (Euphorbiaceael.

species contain

pulmonary complaints and head

Western

those

Equatorial Guinea are Alchornea

infrequently on Senecio alticola and S. erici-rosenii,

ailments.

in

floribunda and

antibacterial,

treating respiratory

utility in

Two other

on Mount Mikeno, DRC."'' Here the animals fed

in

cardiotonic (heart tonic), hallucinogenic,

and stimulatory, or a

'"

hallucinogenic plants eaten by western gorillas and

humans

pharmacological properties

reported

antiviral,

humans

a
it.

jumping around, and fleeing

as though seeing frightening images,'"

118 plant species from 59 families, these plants


being

may have been

watched wild animals ingest

more than one Gabonese source

Information from
of literature

correct, this

If

result of their having

Michael

A.

Huffman and Don Cousins

Western gorilla {Gorilla

(including poachers)
effect,

is

gorilla]

have the opposite

likely to

because larger groups are

lil<ely

be able to

to

be more alert and therefore have a better chance of

defense or escape.'

self

group size ranges from two

Total

32

to

with an average four to six adults,

individuals,

although up to 52 nests have been recorded as


belonging

Box

group"^' " (see the table

to a single

in

Larger groups typically contain a higher

7.A].

proportion of adult females," as most groups are

single-male harems.

Koukoua National

the Maya Nord Bai (Odzala-

In

sometimes as high as nine females per


It

is

assumed

that a female will

her

of

her offspring from predation or

The number

infanticide.

choose whether or

as a breeding partner and

of his fitness

his ability to protect

adult male."

male on the basis

not to stay with a particular

assessment

was

Park], the adult sex ratio

of adult

females

a group

in

would therefore indicate female perception

of the

defensive quality' of the group's male.'"

Upon reaching
gorillas

remain

in

maturity,

the group

(their natal group!, but

some western

in

most emigrate from

females have been observed


groups up

to three times.""'

lowland

which they were born

'^'

it.'"

to transfer

Some

between

Emigrating females

tend to transfer into another group or join a solitary


male.'^'

Male quality

factor

influencing female dispersal patterns.'"'

in

be an important

likely to

is

'''

smaller groups, the foraging costs associated

In

with within-group competition


correlation has been found

the
tive

number of females
success

of

may

a group

in

western

size or

and the reproduc-

gorillas,

although these

results should be treated with caution as the


ple sizes for

(e.g.

at Mbeli Bai in

remain solitary

until they

seven

of eight

in

males

when mother and

group,

but

infant transferred to a

occur on

two

display of a western

occasions.'" Silverbacks gain reproductive benefits

lowland gorilla (here

from

not

which affords a more immediate

infanticide,

nates a potential competitor

to his

Grooming and other forms

own

offspring.

support

of social

are rarely observed between adult females, and

dominance hierarchies also seem

to

female-female relationships being

and ephemeral.'"

'"

Social

be weak, with
individualistic

bonds between western

between mothers and

show

Mbeli Bai

their offspring. Studies in

that offspring suckle for a longer

period than mountain gorillas and remain

can establish their own

proximity to their mothers until weaned. "'"'^ Infants

seem

to

use

this

time

to learn

in

close

about appropriate

foods and

groups form.

niques that are important for an animal that exploits

The mutual dependence between silverbacks


infant protection,

is

gorilla groups.""

When

and

the silverback

in

single-male group dies, the group typically

to

develop the food-processing tech-

such a diverse range

invest
in

developing

members
all

group

time developing

in

these

through

members

as a unit sometimes occurs;

infants.""

new

Infanticide

silverback

sometimes

kills

has been inferred on two

relationships

appear

mountain
with

to

gorillas

group

other

members, perhaps because they are more

disintegrates and the remaining group

circumstances, the

gorillas

less time than Virunga

transfer to

other groups. Transfer of

of habitats.

Young western lowland

impor-

tant in maintaining the integrity of both eastern

western

in captivity).

opportunity to impregnate the mother, and elimi-

between mixed sex groups;"" occasionally, all-male

and females, based on

The chest-thumping

other

did

Congo),"" and generally

Sometimes immature males transfer

group.'"

new

gorillas are not thought typically to be strong, except

males leave the natal group

which they were born


observed

sam-

comparison are small.'"

Virtually all

occasions

No

be lower.'^'

between group

Witliam H Calvin jwwwwiHiamcalvin com]

likely to

leave their natal group. Young gorillas spend

more

can be used later

in life

skills that

play, interacting with

by learning

how

to

respond

from other group members,

younger
to

gorillas,

and

negative behavior

albeit without

forming

117

World Atlas

of Great Apes and their Conservation

A-ta^^izvi

.Ttr~<*

i:

^-

constructed each evening by pulling, bending, and

breaking

stems

the

these

vegetation;

of

are

arranged around and under their bodies." Tree

may be

nests

crown

by folding branches toward the

built

the tree, producing a bed of leaves at the

of

center" Nests may also be at ground

level or

the trees.'"' " Western lowland gorillas

low

in

sometimes

sleep on bare ground, without using vegetation at


all;
1

such sleeping

A5 percent

3 725

of

Hokdu

at

in

more likely to build


" The type of nest

cool or wet weather."

constructed

suggest

thiat,

by the

availability

season, group

materials,

of

size,

microhabitat, and the level of disturbance by other

where an

'''"'

Some western

infant survives, the

animals.

interbirth interval is

nesting

sites

four to six years.

probably

in

Rirridrr)

Karneli

long-term cooperative relationships. An alternative


explanation

access

western gorillas have less

that

is

high-quality food than eastern gorillas

to

have, and therefore

may have

less time to devote to

developing relationships.

The average nesting height varies with the

fifths of

herbaceous undergrowth. Over four

nests are found on the ground at Odzala-

Koukoua National Park


plants like

are

Haumania

common;
the

in

little

nests

swamp

in

found

is

in

the

CAR, where there

in

herbaceous undergrowth.''

is

groups meet often, and more frequently

Congo, where suitable

a similar proportion

Ngotto forest

gorillas do not sleep in

different

in

liebrechtsiana (Marantaceae)

Intergroup encounters

ranges often overlap extensively, so

dense vegetation,

hunting pressure.""

to

Western

gorilla

lowland gorillas select

particularly

in

response

availability of

trees

"

'''

Western

water or on wet ground, so

forests are built quite high up

in

than do mountain gorillas. Several groups often

the trees; at Likouala

make use

example, Raphia (Arecaceael fronds are a favored

of

groups

bais at

same time." Serious

the

between the members

aggression

rarely seen, however; at

is

of

different

in

the northern Congo, for

building material.'^

Lope National

Solitary nests can

be used as an indicator

Park, during 43 intergroup interactions observed

of population health; a disproportionate

from 198A

solitary nests

to 1993, fighting

occasions.'''

one

of

and chestbeats

involve

exchanges

but, in others, only

groups vocalizes. As vocalizations

the

accompany

occurred on only three

Some encounters

of vocalizations

only half of observed encounters,

many

a cause for concern.

is

about 30 percent
percentage rose

to

of

in

Minkebe National

gorilla population

due

to

of the

lead to one group remaining

chapter

in

the area and the

large fruiting trees or groves of trees of

Where

group's range

in

resident'

uncommon

the core of one

had attracted other

silverback of the
to

resource

gorillas,

the

group would display

deter feeding competition.

Response
Western

They usually

from dusk

in

various ways to

human

full

habit-

uation has been reported to produce an increase


in

aggression, fearful reactions, and vocalizations,

Western
gorillas are active only during the
rest

respond

disturbance. Contact with people prior to

as well as longer daily travel distances, but these

Nest building

day.

of

this

to disturbance

gorillas

reactions

Western lowland

in

intergroup encount-

ers were related to access to or defense of either

species.

local

an outbreak

Ebola hemorrhagic fever, discussed later

away Most

of

nests are solitary," but the

60 percent

Park (GabonI following the collapse

western

number

On average,

encounters may go undetected. Many exchanges

other group moving

118

influenced

is

nesting

suitable

Preliminary studies

of

CAR.'"' and

in

Mondika IDzanga-Ndoki

National Park). Gorillas are


nests

Ai percent

sites represented

231 sites observed at Bai

to

dawn

in

nests

diminish as habituation

proceeds.'^'

'

gorillas leave areas to avoid active logging;

they return swiftly unless, as so often occurs, they


are also hunted.

''

Western gorilla [Gorilla

Reproduction

among western

Reproductive development has not been studied

have not yet been conducted across large areas, and

over long periods


birth

at first

two sites

in ttie

western

gorilla,

and

age

ttie

unknown. Preliminary data from

is

Congo, Lossi Gorilla Sanctuary (Odzala-

Koukoua National Park) and Mbeli (NouabaleNdoki National Park), indicate that there are almost
0.2 births per adult

female per year, which

is

to the birth rate of

mountain

has been

gorillas.'

It

similar

areas

lowland gorillas because censuses

swamp

of

identified.

habitat have only recently been

maps

Distribution

western lowland gorillas may remain

gorillas in

swamp

forests, a

variation in food availability could have resulted in

reported only as recently as

the western gorilla having a later age at

be a general pattern

The same

gorilla.

when

the infant survives, the interbirth interval

four to six years.

female
0.62.

reports suggested that

initial

The mean number

Maya Nord

at the

Bai

more than has been seen

in

per

of infants

was reported

is

to

be

any other western

involving

68 females at Lossi Gorilla Sanctuary,


at Mbeli Bail,

widespread habitat that

in

was

989" and confirmed

to

On the other

the 1990s.'^"

hand, the impacts of Ebola and hunting have not

been comprehensively quantified, leading many


researchers

to

caution against overoptimism.'^' The

western lowland

gorilla is classified

indicating that

faces a very high risk of extinction

in

it

as Endangered,

the wild."
Little

and 32 births involving 162 females

of

Congo

previously considered unsuitable for them,

or eastern gorilla group."

Using a very small sample size (12 births

the

many areas numbers may be higher

in

was

and longer interbirth interval than the mountain

numbers

in

than previously thought." The presence of western

hypothesized that increased seasonality and spatial

first birth

often indicate likely

rather than confirmed presence. Large

Basin, '^' and

information

is

gorillas at

1980s,

it

most

sites

in

was thought

numbers,

available on the

status, or trends for populations of

western lowland

most countries.

that there

In

the early

were only 40 000

was ob-

western lowland gorillas worldwide.^' More recent

served to be about 8 percent at Lossi and ^3 percent

estimates have ranged from 9A 500*' to over 1 00 000


individuals;'"' " " several reports have, however,

mortality of infants under one year old

at Mbeli.

The death

rate increased to 22 percent

when

65 percent respectively,

up

to

and

considering infants

indicated that

files in this

screams, hoots, and

been

fully investigated.'^ Gorillas

fruit

trees

call

anticipation of the

decline.

An informal

is

that at

most 82 000

based on mean reported

country figures, adjusted

to

reflect

an estimated

Making charcoal, Congo.

expressions, but the

facial

vocal repertoire of western lowland gorillas has not

often

in

volume suggests

remain. This figure

communicate with grunts, barks,

gorillas

numbers are

and optimistic estimate based on the country pro-

three years of age.'"

Communication
All

gorilla]

Charlie

Semeli-Botarba/UNEP/Topham

running towards

out excitedly,

possibly

in

imminent scramble competition

number

for the

limited

trees. '^'

Hoots are heard more frequently and may

of

feeding spots

in

fruit

be straightforward contact calls to communicate


location

in

the forest.'^"

been conducted
gorillas

No empirical

studies have

to test this hypothesis.

Western

have a larger day range and their foraging

groups are more widely spread than those


eastern gorillas; this

mechanisms

may mean

for maintaining

of

that the vocal

mutual contact are

more important.

POPULATION
Status and trends

Most population surveys

of

western gorillas are

carried out on a site or country basis rather than

with

reference to contiguous populations.

difficult

to

It

is

assess population status and trends

119

World Atlas

of Great Apes and their Conservation

Table 7.2 Western lowland gorilla populations

Date of estimate

Estimated population size

Gabon

1980-1983"

35 000 7 000'

Congo

1990-'"'

34 000-44 000

Cameroon

2000^'

15 000

Central African Republic

1985'

9 000

1989-1990^^

950-2 450

Country

Equatorial Guinea

Democratic Republic

the

of

2000*'

Congo

2000'*

Angola ICabinda province!

From 1983

to

2000

half of all great

apes

loss of half of all Gabon's great

2000.'"

in

Gabon were feared

apes from 1983

Published estimates are sunnmarized

many western

Table 7.2;

'

to

lowland gorillas

be found

Gabon

in

although recent losses are un-

western

both

chimpanzee

and

lowland

gorilla

together,

Gabonese ape populations are thought

to

populations

have declined by more than half between 1983

and 2000.'" A census

in

1989-1990 concluded that

western gorillas were widespread and

common

in

northern Congo. Areas supporting high densities

swamp

included

forests,'

Odzala-Koukoua

of

forests of

and

adjacent

north and

regions to the

outbreak

" such as the


Park

National

" However, an

east.'

Ebola during 2002 and 2003 seriously

affected western gorilla populations at the Lossi

Sanctuary,

Gorilla

southwest
in

of

2004, an

community

itself."

in

likely local extinction is

combined

Riven''

Its

decline and

probably the result of the

effects of habitat loss, fragmentation,

and hunting.'" There

no estimate for western

is

lowland gorilla numbers


of

extreme southwest

the

in

the Cabinda province

Angola.

Threats

Western lowland

gorillas are widely distributed in

a large forested region

numerous protected

and

their

range includes

areas. They nevertheless face

an uncertain future, simply because

western lowland
National Park

of

western gorillas and chimpanzees

Community

is

much

In

CAR

in

ing

scale

and interactive nature

of the increasof

the threats

clearance and road-building, forest degradation by


logging, hunting for food, and disease. Hunting

and

disease are increasing as risk factors because

human access
being

to

formerly remote forest area

is

improved by logging, road building, and

settlement. The estimated halving of the great ape

uncertainty about western


in

1996, western lowland gor-

population

in

combination

Chapter

Gabon

danger

of this

Gabon country

profile,

illustrates the

of factors'" Isee

161.

were assessed as Vulnerable rather than

Endangered
In

120

in

Congo

is

the

clearance for farming, forest fragmentation by

kill

lowland gorilla populations at most locations

illas

of the

in

the park's Lokoue Bai led to

in

eastern Congo."

Cameroon."

DRC, north

gorilla

former range

operating against them.'^ These include forest

the Lac Tele/Likouala-aux-Herbes

There

Bas-Fleuve region
of

its

of

sightings

in

Recent surveys have confirmed still-healthy

Reserve

now absent from

probably

80 percent decline

the Odzala-Koukoua

populations
in

km

The western lowland

industry.'"

oil

'"

fears that Ebola had begun to


in

50

from hunting, forest

to threats

clearance for logging, and/or agriculture and the

Odzala-Koukoua National Park;'"'

western gorillas

gorillas

forest

outside the reserves of

living

Equatorial Guinea have been considered Critically

Endangered due

Considering data on

quantified.

and secondary forest." Western

plantation

of

a national level, the largest populations of

and Congo, '^ "

tost

in

gorilla populations have,

western gorillas are thought

have been

to

however, since suffered declines.

On

to

unknown

Forest clearance, fragmentation, degradation

at a national level.'"

Equatorial Guinea

during

1989-1990, a

Until recently, there

census estimated between 950 and 2 450 western

tat

lowland gorillas to be present," mainly

with

in

areas

has been

degradation over
little

much

relatively little habi-

of

the

Congo

Basin,

conversion to agricultural land."' As

Western gorilla [Gorilla

as the 1980s, West and Central African timber

late

was considered
logging;''"

this

10 percent of

now account

trade recorded

all

in

more than

for

Cameroon, CAR,

Equatorial Guinea, Gabon, and

d'lvoire.

Most

Congo Basin are

of the forests in the

under the control

companies

of

based

that are

in

the European Union, operating either as concession

vationists therefore consider the fate of the forest


wildlife to

its

Europe. By 2000,

be

responsibility shared

more than

by

Gabon's forests

half of

had been allocated as logging concessions," and


production had increased

log

Bushmeat hunting
spread

in Africa,

to

some

is

wide-

In
is

absence

rearing.''" Increasing

the forested

regions of

the main threat to western

of

any tradition

human

of livestock

populations and com-

mercialization of markets have encouraged bush-

meat hunting."

Logging of the western

meat has been reported

Gorilla

food (where available]


of great

apes

for

occurs, for example,


River area

in

in

be a popular

northern Congo. Hunting

in

meat

to

is

widespread

every part

of

in

Congo;

it

the Motaba

the northeast."^ Here, about 5 percent

lowland gorilla's habitat

has increased
dramatically across

its

range since the 1970s,


including here

in

Congo.

3 million

Cameroon, over 170 000 km"

In

for subsistence

Central Africa, hunting

Ian

mVyear.'

the Cabinda province of

both for protein and for sale for

income generation."

Many conser-

holders or as subcontractors."

in

Guinea, Gabon, and Nigeria.""'""""'*'"

gorillas, in the

Liberia."'

and

apes has been reported

Angola, Cameroon, CAR, Congo, DRC, Equatorial

changed dramatically during the

1990s. Forest products

Congo, Cote

low commercial value,

of

pressure posed by selective

the

limited

whicfi

be

to

gorilla]

Redmond/UNESCO

|76

percent] of the country's forests had either been

logged or allocated for logging concessions by then;

images have revealed

satellite

new

logging

roads have

that

networks

now spread

of

what

into

had previously been considered the least accessible forests in the country."'

range

of the

western lowland

Other parts

gorilla to

of the

have under-

gone extensive logging include the mainland

of

and the Congolese sections

Equatorial Guinea

Mayombe forest." '"'^'Although logging ocsome of the protected areas in Cameroon,


CAR, Gabon, and Congo that are home to western
lowland gorillas,'"""'" many others have escaped

of the

curs

in

intact so far.

Logging roads and access routes fragment


as well as improving access for hunters.

forest,

Forest fragmentation poses a potential threat to

western lowland
to

groups.

gorillas, in that

it

can block access

sources and prevent transfer between

food

It

is

unclear what degree of fragmentation

constitutes a barrier to western gorillas,'^" but

the continuous forest and

National Park

savanna

Gabon, western gorillas were

in

reported to be reluctant to cross gaps


that

in

the Lope

of

were wider than 50

the forest

in

m.'^'

Hunting

Western lowland gorillas are hunted


for

sale

to

private

infants), for trophies,


ritual
to

the

collections

and

for traditional

purposes. Although this


national

laws

of

for their

is

medical or

illegal

and

according

every range state, the

regulations are often poorly enforced at


of the legal

meat,

(particularly as

judicial system.'^

all levels

Hunting

of

great

121

World Atlas

of Great Apes and their Conservation

of

the western

estimated

lowland gorilla

despite the sparse local


level of hunting

mammals
The
gorillas

is

was

human

population. This

unsustainable for slow-breeding

western lowland

varies throughout their range.

legislation

in

many areas

of

respectively.'''"

West-Central Africa,

gorillas are killed or injured by snares set for other

species."

Factors

bushmeat hunting are

local

enforcement],

the

land

its

ammunition and guns, and the ease

of

much

threats over

Cameroon. Hunting

habitat of
to the

small population

is

a particular threat

River. Although

Reserve, hunting does occur, and timber extraction

remote areas and by bringing a hungry workforce


the forests." Western lowland gorilla popu-

lations have declined


Civil

where timber extraction has

wars

in

DRC and

civil

unrest

Congo and CAR have also increased hunting

in

levels

by exacerbating poverty and dependence on wild

resources, particularly

among

western

to
in

was

between September 1966 and February 1969, many


destined

for

centers.'"

The capture

due

to

"

well known." As early as the 1980s,


believed

to

of

and

parks

zoological

and export has declined due

for sale

research

western lowland gorillas

and

to national

Angola, western lowland gorillas have been hunted

although the negative impact of hunting on gorilla

hunting

western

the overall

of either

gorilla population losses specifically

is

live

international conservation efforts, but live infants

hunting, or of their impact on population trends,"

populations

the areas surrounding the reserve.'"

lowland gorillas were removed from the wild

are

There are no estimates

in

Equatorial Guinea, at least 63

In

displaced peoples

and refugees.

be the primary threat

western lowland gorillas and chimpanzees


Gabon, as their population density was lower

sometimes traded

still

border"" This

is

trade,

and

live

Luanda and across the

infants sold in the capital

which has

within the region.'^" In

bushmeat

recent years for the

in

largely a result of the conflict there,

immigration

led to the

taboos on eating apes moving

of

people without

into

areas where

these taboos have traditionally operated.

in

contrast, effective controls on hunting are

In

areas where the animals were hunted, with density

in

reductions of 17 percent and 72 percent observed

buffer zone ICongol,

Kelley McFarland

place

Nouabale-Ndoki National Park and

in

Park ICongol, and

in

in

its

Odzala-Koukoua National

the

Dzanga Sector

of

the

Dzanga-Ndoki National Park ICAR), where apes


are rarely hunted."'" The controls are the result

gunshot hole and

of

embedded

governments and outside agencies: the

(indicated by arrows).

no

has taken place within the Dja Faunal

logging

continues

occurred."

lead shot

be

to

gorilla

western lowland gorillas

of

Sanaga

the north of the

to

western lowland

of the

roads contribute both by promoting greater access

into

gorilla skeleton with

and heavy hunting

of light

hunting under local seasonal climates." Logging

to

River

areas

Furthermore,

Hunting and logging are considered

intensity of hunting of

availability of

A female Cross

in

like gorillas."

affecting the intensity of

taboos,

population

be killed by hunters each year."

to

successful collaborations between the national

Conservation

Society

Wildlife

Nouabale-Ndoki;

in

the

European Union program. Conservation and Rational

Use

Forest Ecosystems

of

lECOFACl

in

in

Central Africa

Odzala-Koukoua; and

WWF-The

Global Conservation Organization, along with the

German overseas development agency GTZ,

in

Dzanga-Ndoki National Park.

Disease
Disease

is

a potentially devastating threat to great

apes. Western gorillas are susceptible to


of the

same

virus,"

'^"

chicken

the

pox,

many

diseases as humans, such as Ebola

common
bacterial

measles, rubella,

cold,

pneumonia, smallpox,

meningitis,

mumps,

tuberculosis,

yellow fever, encephalo-

myocarditis, and paralytic poliomyelitis"'" Of the


identified

122

western lowland gorillas

in

the population

Western gorilla [Gorilla

at

Maya Nord

gorilla]

example, 5.7 percent were

Bai, for

reported to be affected by yaws (frambesia tropical,


while

some

others showed signs

of

the onset of this

disease." There are several varieties


the pathogen

gorillas

in

yaws, but

of

probably Treponema

is

pertenue.^' The disease causes tissue necrosis;

common

humans

in

it

is

Central Africa and can be

In

treated with antibiotics.

has been assumed that already small or

It

fragmented populations are most vulnerable

otherwise. Ebola

human

is

disease that

to

shown

disease," but Ebola hemorrhagic fever has


best

known as an Incurable

kills

about 80 percent

of Its

victims. This virus has an even higher mortality rate


of

95-99 percent among chimpanzees and western


Recent Ebola epidemics

gorillas.

West

in

have affected the western lowland gorilla

to

have

great

ape

and Congo. Ebola outbreaks are thought


contributed

strongly

populations

known

to

in

decline

to

of

Africa

Gabon

in

Gabon, where four outbreaks are

have occurred, two

of

which originated

in

the Minkebe National Park." Farther east, declines


In

western

gorilla populations attributed to

have also been


Sanctuary

of

reported

in

the

Lossi

Ebola

Gorilla

may

Congo, with fears that the disease

However,

It

seems

that

It

was Ebola

disease not

la

associated with tourismi that killed eight groups


of

habituated western gorillas at Lossi

Gorilla

Sanctuary between October 2002 and January


2003."

human outbreaks

of the recent

western equatorial Africa appear


Initiated

when people handled

the

in

to

have been

meat

of infected

great apes.
Habituation, the process

come

whereby apes be-

tolerant of the presence of

humans, allows

regular and consistent observations by researchers

and by

tourists.

proved

difficult

dense vegetation

The western lowland


to

particularly as

habituate,

does not allow

of Its habitat

tracked easily.'" Gorilla tourism

as well established as

it

Is

gorilla

is

It

for eastern gorillas.

Habitat protection and law enforcement

Western

gorillas are legally protected from per-

secution

In

all

range states. There are protected

areas within the range


species, but

Due

to

most

the sparse

of both

human

population

The

protected,

many areas

although this

is

In

sub-

much

of its

changing rapidly as
Its

toll.

The

protected areas that host western lowland gorillas


include a World Heritage Site

Faunal Reserve, an area

like-

that are not formally

logging spreads and hunting takes

be seen

however, has increased the

gorilla

relatively well in

discovery that western lowland gorillas could easily


at bais,

western

gorillas live outside these.'""

the

therefore not

of

in

Cameroon

(the Dja

260 km^l, various

lihood of successful gorilla tourism. Tourism can

national parks including Dzanga-Ndoki (CARI, Lope

provide significant revenue that can be channeled

(Gabon!, Monte Alen lEquatorlal Guinea], Odzala-

into

ape conservation, but

number

of

people

in

it

also

increases the

daily contact with gorillas. This

Koukoua,

and

Nouabale-Ndokl

(Congo),

increases the chance of disease transmission, and

River gorilla occurs

Cross River National Park (Nigeria; see Box

with

humans and undergoing

their resistance to disease."'

habituation

contact

may lower

Although regulations

and

several other categories of reserve. The Cross

the stress experienced by gorillas while

In

about the disease risks

great apes.

range, the western lowland gorilla has so far fared

be

in

villagers

CONSERVATION AND RESEARCH

has

to

Congo teach

of contact with

have spread Into the Odzala-Koukoua National

Park.^"'" Many

Village outreach:

education efforts

The
afforded

level

of

in

several reserves including


7.1

1.

nominal and actual protection

by the different protection categories

help protect eastern gorillas from disease trans-

varies between countries, reflecting their different

mission from tourists and their guides," no such

histories

regulations yet exist for western gorillas. Guidelines

range states are resources for conservation abun-

have been developed for Mbeli Bal and Bai Hokdu.

dant, however; all the countries Involved are

and economies. Nowhere

in

the gorilla

among

123

World Atlas

of Great Apes and their Conservation

Box

lA GORILLA CENSUSES

reasons.

on

most western

In

the

most

method

number and

density.

A common method

number

of nests

seen per kilometer

the

between

estimating gorilla

of

to

is

in their durability

is

differences

climatic

can therefore influence the results.

sites

The nest-decay

count

of transect

depending

rate

used

in

population calculations

intended to correct for this."^'

In

the absence of

walked. The size of the associated gorilla groups

further data, manystudiesrelyon the nest-decay rate

can be estimated by examining

of Tutin

nests at a nest-

source

gorillas are

found at a density of about 0.25 weaned

typically

individuals per square


this

all

Western lowland

ing site Isee table).

many

can be as

gorillas per

some

kilometer At

as three

between forests

differ

such

resources

and

National Park, for example, the

are

further complicated

methodology and

The number

and established

by differences

of

weaned

individuals

of nest counts,

in

an area

group

size

is

sizes,

nest-decay rates Ithe rate at

local

in

of

foods.

western gorillas

number

number

of gorilla

weaned

of

the group at only one third of fresh nest

Large sample sizes are required

At bsis.

field

in

same

to

minimize

the impact of these possible errors.'^'

directly

disintegrate], often using a

transect-analysis

sites-'^^

level of effort."

estimated on the basis

which nests

individuals

population density

of

of the

gorillas sleep on bare ground,

nests corresponded to the

'" "'

herbaceous plants."'

as

comparisons

Interpretation of

and eat many

present can be underestimated.^'''"'' "'''" At Lope

the abundance and distribution of key

in

trees,

which means that the number

species composition, and are affected by subtle


variation

in

some western

Third,

may host

similar structure

of

uncertainty Second, western gorilla and

often nest

sites,

or, exceptionally, five

square kilometer; poor habitat

and Fernandez,'" introducing an additional


of

chimpanzee nests can be confused, as both species

as fewasO.l/kml Western lowland gorilla densities

computerized

It

IS

possible to observe western gorillas

and obtain very accurate information on the

and composition

size of

groups seen

corded

in

of

groups. Typically, the

at bsis is larger

mean

than that re-

nest counts elsewhere. However, these

findings relate only to those western gorilla groups

program, such as DISTANCE.^'

with

access

to

these clearings; supplementary

Nests can also provide information on the age

research

structure and gender balance of populations, but

density and the distance traveled to reach a bai.

must be analyzed with care

their results

Group

size in

km

of

western lowland gorillas

transects

Dzanga-Sangha
Ngotto

forest,

Number

in

region,

CAR^' 194

CAR^' 1783

km

km

swamps, Congo'^

swamps. Congo"'"

180

km

1401

of transects,

km

wet season!

of transects,

dry season!

Bai Hokou, CAR'^ lobservation and nest counts!

Maya Nord

Bai,

lobservation from viewing platform!

Odzala-Koukoua National Park, Congo

lobservation from viewing platform!

a Data are expressed as

mean

plus range, where available,

b Including 73 solitary males.


c

Where data

are not available, this

is

indicated by a dash.

d Excluding solitary males.


e This group, unusually, had two silverbacks.
f

Number

of

Weaned

gorillas

nests

per group"

136

540

4 11-191

261

of transects!

of transects!

Likouala

Congo

of

groups

15 habitat types!

Likouala

Mbeli Bai,

required to discover the local population

Sarah Fernss and Lera Miles

Northwestern Gabon""
1782.8

is

for several

Location and study method

124

nests vary

First,

they are built and the weather to which they

are exposed; vegetation and

counts are

gorilla habitats, nest

practical

how

The range observed over a 27 month

period.

323

5.1

11-521'

_'

145

5.712-111'
5.6 12-10!"

38

213

36

5"

12-15'

14

8.4 4.3"

36

11.2 12-221"

Western gorilla [Gorilla

the world's poorest. This puts a

premium on

reliev-

legislation

and the degree

enforcement. Efforts

of

ing constraints on conservation resources through

made

partnerships between range state governments,

areas, especially those that straddle frontiers, and

donor agencies, and nongovernmental

official

organizations. For example, since the gazetting of

Dense Forest Reserve

the Dzanga-Sangha Special

by

many range states in

establishing protected

the protection of western gorillas

show

official

commitment

gorilla habitats. In all

to

in

national law,

the conservation of

range states, however, lack

resources and financial constraints impede

been managed by the the Dzanga-Sangha

efforts to enforce existing legislation effectively.

collaboration

GTZ

between the CAR government,

German

(acting through the

LUSO

WWF"

and

Consult),

new law

consulting firm

Partnerships with the

Congo

private sector can also help;

innovative

project,

which

into place,

putting an

is

will require

logging concessions to provide patrols to dis-

all

courage poaching.'"

Some
protected

Conservation and research activities

The large range

of

the western lowland gorilla

encourages some confidence


yet

brings

its

in its

own challenges

survival chances,
of

coordination

between the multiple governments and other


stakeholders.

blocks of western gorilla

Many

international,

regional,

and

or

national organizations are working to safeguard the

areas straddle international borders,

western lowland gorillas future through conser-

requiring cooperation between two or


tries for effective conservation.

habitat

Forested mountains
at the

of

headwaters

Asache

River, a

Cross River gorilla


survey area, and a
survey

camp
same

field

in

the

area.

more coun-

One such cross-

border regional collaboration has been

mented

of

many

and the Dzanga-Ndoki National Park, both have

gorilla]

imple-

southern CAR, northern Congo, and

in

southeast Cameroon, establishing Trinationale de

Sangha

la

in

This conservation

1998.

initiative

covers the contiguous Dzanga-Ndoki. NouabaleNdoki, and

Lobeke National Parks, and divides

the area into regions

managed

which human

in

or restricted.

It

activity is

allows for joint patrols by

rangers from the three countries and has resulted

some

in

successful missions

in

the ongoing effort to

discourage poaching."

Although parts
protected

in

much degraded.
underway
forest

the

of

has been

proposed system

The Mengame

to relieve

transfrontier

three countries.'"

all

Sanctuary comprises

Gorilla

000 km'' biodiversity corridor


border with Gabon.

of

is

promote

to

and biodiversity conservation, and

protected areas involving

its

forest are
it

Dialog with local communities

Cabinda province, aiming

in

poverty via

Mayombe

Angola, DRC, and Congo,

It

in

will

Cameroon, along
contribute

to

transborder protected area by linking with the

Minkebe National Park


emerging tri-national
(Dja

in

Gabon, as well as

initiative

to

an

between Cameroon

Faunal Reserve), Gabon (Minkebe National

Park),

and Congo (Odzala-Koukoua National Park).


Jane Goodall

Institutes

signed an

agreement with the Cameroon Ministry

of Environ-

In

2002,

the

ment and Forests (MINEF)


based conservation and
the

to establish a

wildlife

Mengame Gorilla Sanctuary.^"


A key constraint on the success

gorilla

conservation

is

community-

research program

of

in

western

the quality of protective

125

World Atlas

of Great Apes and their Conservation

programs (more than can be

vation and research

mentioned
provide

some

active in

each

of the

the

from human disease are often poorly enforced,

with

the most sustained initiatives

of

GoriUes

des

was

established by Tutin and Fernandez with

tial

funding from the Centre

due

mainly

des

et

lack

to

of

using hides or observation platforms, from which


tourists could observe the wildlife without the

ini-

International de

to

Recherches Medicates de Franceville ICIRMFI, and

contact with them. Experience from zoos has taught

us that gorillas become anxious

studies of western lowland gorilla and chimpanzee

from above,'" so where

have been conducted there since 1983. The regional

hide would be worth constructing.

of

ecotourism

assisted

the development

in

supporting ecological and sociological studies


area.

ECOFAC was

initiated to

areas and the development

parts

the

of

ECOFAC has undertaken

of

investment

of

and

representing a significant

habitat,

its

resources and the

scarce public

protected

setting aside of large areas of forest land. Threats

sustainable local

are nevertheless increasing, and originate largely

of

throughout Central Africa.' Since 1992,

activities

when watched
might be used, a

gorilla are taking action to protect the species

ensure biodiversity

management

conservation through

in

a platform

conclusion, the range states of the western

In

Lope National Park, as well as

at

need

habituate the gorillas or for there to be any

with later support from other donors. Long-term

ECOFAC program has

and

training

staff

education.'" Viewing at bais offers the option of

Lope National Park, Gabon, which

in

However,

rules designed to regulate tourism and protect apes

hand-in-hand

goes

d'Etudes

Station

Chlmpanzes

be channeled into ape conservation.

experience with the eastern gorilla has shown that

range states.

often

One

conservation.

Tourism can generate significant revenue that can

Chapter 16

profiles in

further details about organizations

Research

is

The country

here].

in

the unplanned and unregulated infrastructure

development

surveys,

biodiversity

associated

with

particularly

the

including the collection of information on primate

timber industry. This renders very large areas

as Odzala-Koukoua

accessible to hunters and encourages development

populations at sites such


National Park,

Faunal Reserve, Monte Alen

Dja

also

further endangering

have also played their part

been established. Attempts

already vulnerable western gorilla populations. Our

to the wild are at

Projet

gorillas

number

of

an early stage.

In

Congo,

collective

Reserve.^''

these pressures

mitigate

to

ability

in

limited by ignorance of both the status

among
much

orphaned western low-

Lesio-Louna

the

into

reintroduce

to

des GoriUes successfully

Protection

reintroduced a
land

including outbreaks of the Ebola virus,

disease,

Sanctuaries for captive western gorillas have

orphans
the

bushmeat market. Habitat fragmentation and

of the

National Park, and Ngotto forest.

"

populations
of

the western gorilla across

of

research

Further

range.

its

is

and trends

on

the

distribution,

abundance, and status

one sanctuary that accepts orphaned western

gorillas

their

lowland gorillas; Angola, CAR, and

needed, with a focus on the Cross River gorilla

Cameroon,

Guinea do

Nigeria,

and Gabon also have

Equatorial

being

not.

There are several efforts

to establish

operations based around sightings

of

salt clearings [bais] just north of

Park,

western lowland

visited

matter

behavior would

western

range states

of

of

is

western

therefore urgently

particular

western

of

priority.

gorilla

Better

ecology and

improve the likelihood

of

con-

servation success. Long-term studies would be the

Odzala-Koukoua

which are regularly

in all

understanding

tourism

Congo, Cameroon, Gabon, and CAR. The

gorillas, in

National

at least

ideal

way

to

meet both needs, while also being

proven way to promote conservation

by

in

and around

the study sites.

gorillas, are potentially suitable.

FURTHER READING
Cipolletta, C. 12003)

Ranging patterns

of a

western

gorilla

group during habituation

to

humans

in

the Dzanga-Ndoki

National Park, Central African Republic. International Journal of Primatotogy2i. 1207-1226.

Cousins.

D.,

Huffman, M.A. 120021 Medicinal properties

in

the diet of gorillas: an ethnopharmacolcgical evaluation.

African Study Monograpfis 23: 65-89.

Doran, D.M., McNeilage,


resource

American Journal

126

A.,

availability:

Greer,

new

D.,

Bocian, C, Mehlman,

P.,

Shah, N. (2002) Western lowland gorilla diet and

evidence, cross-site comparisons, and reflections on indirect sampling methods.

of Primatology58: 91-116.

Western gorilla {Gorilla

Harcourt, A.H. 119861 Gorilla conservation: anatomy

Harcourt, A.H. (19961

campaign.

of a

New

Self-sustaining Populations. Springer-Verlag,

Primates: The Road

K,, ed.,

to

York. pp. 31-46.

How

the gorilla a threatened species'

Is

Benirschke,

In:

gorilla]

should

we judge?

Biological Conservation 75

121:

165-176.
Iwu, M.M. (1993)

Magtiocca,

Handbook

Querouil,

F..

S.,

of African Medicinal Plants.

CRC

American Journal

gorillas in north-western Republic of Congo.

Nowell, A.A., Fletcher, A.W. (20041 Behavioral development

mountain

gorillas. Folia

Parnell, R.J. (2002)

Press, London,

Gautier-Hion, A. (1999) Population structure and group composition of western lowland

Group

in

of Primatology i8 HI: 1-14.

comparison with

wild western lowland gorillas and a

Primatotogica 75 SI: 314.

and structure

size

in

western lowland gorillas

{Gorilla gorilla gorilla) at Mbeli Bai,

Republic of Congo. American Journal of Primatology56: 193-206.

Buchanan-Smith,

Parnell, R.J.,

H.l^. (2001)

An unusual

Peterson, D. (2003) Eating Apes. California Studies

in

Robbins, M.M., Bermejo, W., Cipolletta, C, Magliocca,


history patterns

in

western gorillas

Rogers, M.E., Abernethy,


six sites.

Sarmiento,

F.,

6.

Nature UM: 294.

University of California Press, Berkeley

Parnell, R.J., Stokes, E. 12004) Social structure and

[Gorilla gorilla gorilla).

Bermejo, M., Fernandez, M., Tutin, C.E.G. (2004) Western

K.,

American Journal

Pnmatology

of

gorilla diet: a synthesis

from

173-192.

bit:

The Cross River

E.E., Dates, J.F. (2000)

life

American Journal of Pnmatology6i: 145-159.

gorillas: a distinct subspecies. Gorilla gorilla diehli

Matschie

American Museum Novitates 3304: 1-55.

1904.

Stokes,

social display by gorillas.

Food and Culture

E.J.,

Parnell, R.J., Olejniczak, C. (20031

Female dispersal and reproductive success

gorillas [Gorilla gorilla gorilla). Behavioural Ecology

Taylor, A.B., Goldsmith, M.L., eds 120021 Gorilla Biology:

in

wild western lowland

and Sociobiology 54: 329-339.


A

Multidisciplinary Perspective.

Cambridge University

Press, Cambridge, UK.


Tutin, C.E.G.

Fernandez, M. 11984) Nationwide census

troglodytes) populations

of gorilla

[Gorilla g. gorilla)

Gabon. American Journal of Pnmatology

in

6:

and chimpanzee [Pan

t.

313-336.

MAP SOURCES
Map

7.1

Chapter

Gorilla data are

based on the following source, with updates as cited

Butynski,T.M. (20011 Africa's great apes.


A.,

in

the relevant country profiles

in

16:

Stevens,

E.F..

Arluke,

A.,

In:

eds, Great

Beck, B.B., Stoinski,

Hutchins. M., Maple,

T.S.,

Apes and Humans: The Ethics

T.L.,

Norton,

B.,

Rowan,

of Coexistence. Smithsonian Institution

Press. Washington, DC. pp. 3-56.

Box

7.1

Cross River gorilla data are based on unpublished data from Richard Bergl and Jacqueline

Groves, with additional data as cited

in

the

Cameroon and

L.

Sunderland-

Nigeria country profiles.

For protected area and other data see 'Using the maps'.

ACKNOWLEDGMENTS
Many thanks

to Kelley

McFarland

Alexander Harcourt (University


City University of

Jacqueline

L.

their valuable

on

gorilla

New

(City University of

of California, Davis),

New

York) for information on Cross River gorilla diet,

Michael Huffman (Kyoto University), John

York), Richard Parnell (University of Stirling),

Sunderland-Groves

comments on

(Wildlife

Emma

F.

and

to

Gates (Hunter College,

Stokes (Wildlife Conservation Society),

Conservation Society], and Elizabeth A. Williamson (University of

Stirling) for

the draft of this chapter, and to Angela Nowell (University College Chester) for information

development. Thanks also

to

Muhammad

Akhlas (UNEP-WCMC) for research

into the literature.

AUTHORS
Sarah Ferriss.

Box

7.1

UNEP World

Jacqueline

L.

Conservation Monitoring Centre

Sunderland-Groves, Wildlife Conservation Society, John

Society and Hunter College, City University of

New

York,

F.

Gates, Wildlife Conservation

and Richard Bergl, Hunter College,

City University of

New York
Box

7.2 Richard Parnell, Scottish

Box

7.3

Primate Research Group, University

Michael A. Huffman and Don Cousins, Primate Research

Box 7.4 Sarah Ferriss and Lera Miles,

UNEP World

of Stirling

Institute,

Kyoto University

Conservation Monitoring Centre

127

World Atlas

of Great Apes and their Conservation

Martha

Robbins

Eastern gorilla {Gorilla beringei]

Chapter 8

Eastern gorilla
[Gorilla beringei]
Sarah Ferriss, Martha M. Robbins, and Elizabeth

Eastern

[Gorilla

gorillas

19031, occur

in

the wild

beringei Matschie,

There

km

differences

more than

from the nearest western gorillas

000

[6. gorilla

Savage, 18A7I. They are larger than the western


gorilla but

otherwise similar, with a broad chest and

shoulders, a large head, and a hairless, shiny black


face.

220

full-grown adult male can weigh up to about

kg,

and a full-grown adult female about

" " Two

this."'

subspecies

currently recognized

Group

of

of

by the

half

Union:'" the
b.

16.

graueri
16.

b.

in

gorilla

hair,

and

tends

occurs

Mount

on

of

the

Congo

(DRC)."^

population of mountain gorillas, that found

Impenetrable National Park

distinguished by

is

less

rounded and more angular nostrils." Genetic

in

in

One
the

Uganda,

confirmed by comparison

their mitochondrial

of

ImtDNAl;" however, the high frequency

recently reported

in

gorillas'"'"'

tion of the earlier results


of

these two subspecies

more

viously

been thought.

It

tentious issue, as

short

is

it

gorilla populations

turn out to be

estimated that the two

DISTRIBUTION
Mountain

gorilla

The mountain

probable that the mountain

have been separated for only the

period

during which

intensive

Uganda [Map

gorilla

occurs

gorillas

are

Rwanda

Mgahinga

and the small number

make

of

more

it

gorilla populations

samples available
difficult

for

than usual to

determine whether the variation between popugreater than the variation within them.

Debate continues on

this issue,"'

""""^

recognize only two subspecies

the

mountain

eastern

gorilla.

lowland

of

but here

the eastern

gorilla

and

the

1.

DRC, Rwanda, and

These populations occur almost

One

is

found

among

protected

officially

by the Virunga

National Park of DRC, the Volcanoes National Park


in

mountain

size of the

8.1

two known popu-

in

the extinct volcanoes of the Virunga Massif. These

agriculture has occupied the area between them.

gorilla:

more

than had pre-

different)

subspecies diverged some 400 000 years ago."'

The small

we

interpreta-

may
is

of

DNA

The mtDNA

difficult.

entirely within national parks.

is

makes

more

some have suggested that it should be


considered a third subspecies." '" This is a conhavior that

lations

was apparently

divergence of these subspecies

lations within three countries:

examination

its

larger cranium and wider facial skeleton, as well as

has such distinctive morphology, ecology, and be-

relatively

al-

have a larger

to

the Virunga National Park of the

Democratic Republic

Bwindi

body and longer

similar (or indeed

very small population of unusually large

Tshiaberimu,

though the mountain

incorporation of mitochondrial into nuclear

Matschie, 19141; and the mountain gorilla

gorillas

physical

ONA

lUCN-The World Conservation

lowland

absolute

Primate Specialist

faer/nge/ Matschie, 19031.

any

if

between these two subspecies,

eastern gorilla are

eastern lowland or Grauer's gorilla

eastern

few

are

Williamson

A.

IParc National des VolcansI, and the


Gorilla National

Park

in

Uganda;

all of

these are contiguous and so protect a single area of


gorilla habitat (the Virungasl.
is

found mainly

Park

in

in

The other population

Bwindi Impenetrable National

southwest Uganda on the border with DRC.

Mountain gorillas occupy about 375 km'


Virungas and 215 km'

in

in

the

Bwindi," these areas being

separated from each other by 25

km

of settled

farmland."^

129

World Atlas

Map

130

8.1

of Great Apes and their Conservation

Eastern gorilla distribution

P^la sources are provided at the end of

this

chapter

Eastern gorilla [Gorilla beringei]

The three national parks

most

contain

UQ kml

extending to about

ranges

elevation from 2 000

in

the Virungas

of

of the surviving forests in the region,

montane

occupy an estimated 15 000 km'

Park and adjacent

and bamboo

tree cover

little

and

forest; the

Itombwe

Forest; and

North Kivu."
Kahuzi-Biega covers an area
ranging

altitude

in

from 600

to

6 000

of

park

1600 km^l and a lowland sector

combine

nected by a forested corridor Gorillas occur

marshes, swamps, and

is

km',

3 400 m. The

high rainfall, and complex topography and drainage


lakes,

four

the adjacent Kasese region; the Maiko National

4 500 m, and

to

abundant herbaceous vegetation, The area has a

to create

in

rain forest formations

stands, as well as areas with

to

broad regions: the Kahuzi-Biega National Park and

The forested area

therefore contains a range of ecosystems, including

various

thought

divided into two parts, a mountain sector


(5

400 km'], conin

both,

peat bogs at various altitudes. The volcanic history

in

means

that there are high eroding

the mountain sector, and the Kasese region of the

plains,

and that

generally high; this

soil fertility is

principally responsible for the high density

factor

is

of the

surrounding

human

eastern DRC,

km

is

almost 800

km^

7 900

length.

in

area,

in

located

is

north-

and has a bound-

ranges

It

in

m, so contains non-

to over 5 100

as few as

located

in

DRC

percent of the park

west side

North Kivu iKivu NordI

in

Province and the remainder

is in

Orientate (formerly

The park has four sectors, with

Haut-Zairel.'^'

gorillas found only

in

The Volcanoes National Park


about 160 km^
2 400

15

area and ranges

in

m to 4 507 m.-

km

northwest

'^'

Its

boundary

town

of the

of

is

in

area, ranges

4 127 m, and

Gorilla National

altitude

in

located

is

in

is

altitude

from

located

some

Ruhengeri

DRC

Virunga Massif on the Ugandan and

The Mgahinga

Rwanda

of

in

Park

in

the

about
to

and the mountain ranges

300

of the Rift Valley.

800 km" and ranges


m."-*

''
It

is

33.7

from around 2 400

km^

around the area

unclear

in

altitude

how

of

from 700

war

in

heavily the

Maiko's gorilla population. There

of the Maiko, Virunga,

and Kahuzi-

Biega National Parks; together, these are thought


to

host between 700 and

number

of

400 gorillas as well as

chimpanzees.

'^

One

of

these encompasses the Itombwe Forest, an area

to

of

to

DRC and Rwanda. The

is

of the

The park has an area

are also several developing community reserves

the extreme southwest of

Uganda, on the borders with

the upland region between the central

river basin

an unknown

borders.

from an

000 individuals."^

DRC has impacted

the southern sector

late 1990s, falling

an unknown number, perhaps

to

Maiko National Park and nearby forests are

forested areas at low and high elevation. About 95


is

in

Gorilla populations in this park

war during the

civil

estimated 8 000

from

altitude

in

'^^

lowland sector"

Mount Kahuzi

are thought to have been devastated during the

DRC

population.

The Virunga National Park


ary 650

peaks and lava

the region of Lake Kivu and

montane,

transitional,

and lowland tropical forest

the west of Lake Tanganyika,'"' and includes

Habitat of the eastern


tovifland gorilla,

Tayna

Gorilla Reserve,

Democratic Republic

of

the Congo.

Pierre Kakule Vwirasihikya

park was established specifically for the protection


of

mountain gorillas

1991, having been a desig-

in

nated but unprotected reserve prior to

The second population


is

mainly found

located

in

in

the

DRC

mountain

of

" " ''

gorillas

Bwindi National Park, which

Highlands

Kigezi

Uganda, on the edge


bordering

that.'^-

of the

Albertine

is

southwest

of

Rift Valley

and

Some of these gorillas


border in DRC itself. Bwindi

to the west.'""'

also occur across the

National Park covers approximately 331 km^ and

ranges

in

altitude

from

160

Mountain gorillas occur

to 2

607 m.

at a density of 0.85-

1.00/kmMn both Bwindi and the Virungas."' ''"''"


Eastern lowland gorilla

The eastern lowland

gorilla

occurs only

in

eastern

DRC, between the Lualaba River and the Burundi-

Rwanda-Uganda border

Its

ses an area of about 90 000

distribution

km^

encompas-

within which

it

is

131

World Atlas

of Great Apes and their Conservation

as high as U 100 m, while those

to

^K^^^^

rT^

live

r
I

between

occur

gorillas

"^
9m.

160

JGkr^
-

J^K^jj

their populations are geographically separated.

The Virungas mountain

^-

i*S

mountainous volcanic

number

region that contains a

The most widespread type

of vegetation zones.

Hagenia abyssinica

is

IRosaceael and Hypericum revolutum (Clusiaceael

.^

."

.:

..

grassy understory.by mountain gorillas

found

Park.

reserve
in

000 l<m^

in

is

also being developed

Nortfi

Kivu area, to

tfie

less frequently,

Other areas frequented

the Virungas include open

excellens (Acanthaceael,
Gordon Mdler/IRF

It

wfiich gorillas are

in

in

""

in

the

flat

saddle between

Mounts Visoke and Sabinyo; monospecific stands

A community

four separate populations."

in

ttie

11

''

or,

herbaceous areas, often dominated by Mimulopsis

protected areas of several different designations.

covers an area of

Impenetrable National

open canopy and

relatively

extremely dense herbaceous

"v-

Bwindi

gorilla habitat

a forested,

woodland, with a

Habitat of the mountain

and

the eastern lowland and mountain gorillas overlap,


L_

^Ke

gorilla,

between 600

altitudes

at

Bwindi

at

600 m. Eastern lowland

2 900 m."'''''-'' Although the altitudinal ranges of

The Virungas are

^K.

and

of

bamboo; dense ridge vegetation with abundant

Hypericum revolutum and shrubby growth

of

Senecio mariettae (Asteraceael; and high-altitude

m.

vegetation with a stature of i-5

the Masisi region

southwest

of the

Virunga National Park.

Bwindi mountain gorilla habitat


Bwindi gorillas

live at

lower elevations, and are more

arboreal than gorillas of the Virungas."^ They occur

BEHAVIOR AND ECOLOGY


Much of the information on the ecology of the
eastern gorilla comes from studies on mountain
gorillas

in

the Virunga Massif."- -

the Virungas
in

was

Research

by George

initiated

graphy

of

trees and shrubs, usually interspersed with lianas


riverine

along permanent or temporary rivers or

streams, with an open or continuous canopy; and regenerating forest that has been disturbed previously,

in

in

Bwindi," or of the

but research

for

example by logging." There

ongoing.

fruit-bearing trees

in

eastern lowland gorilla have

National Park than

in

gorilla,

of the

been carried out

is

is

a greater density of

the gorilla habitats of Bwindi

the Virungas.^"'

Kahuzi-Biega, under the aus-

pices of bodies such as the Congolese Institute

Nature Conservation (ICCN, Kinshasa), the

Center

forest,

in

eastern lowland

for

and woody vines, especially Mimulopsis spp.;

known about the ecology or demo-

mountain gorillas

Most studies

by Mimulopsis arborescens (Acanthaceael;

Schaller

researchers from the Karisoke Research Center


is

inated

open

sometimes dom-

mixed forest dominated by understory and canopy

groups have been followed regularly by

Rwanda. Less

a range of vegetation types which include

in

the late 1950s; since 1967, three to four habit-

uated

in

forest with a discontinuous canopy,

Eastern lowland gorilla habitat

The eastern lowland

Natural Science Research ICRSN, Lwirol,


and Kyoto University.^'' " '" '"' Research findings

dinal

on the ecological and behavioral differences be-

tropical forests.

tween and within the eastern

of

of

gorilla subspecies,

and between western and eastern

gorillas,

are

accumulating.'"'-''"''''''"'-'"^

gorillas, living in

densities:

0.55/km'
in

gorilla

has the widest

and geographical range

at

montane,

of

any

of the

transitional,

and lowland

They have been reported

0.25/km'

in

altitu-

eastern

at a

range

Maiko National Park,

Mount Tshiabehmu, and 1.03-1.26/km'

Kahuzi-Biega.""""

'"

One

of the best-studied

populations of eastern lowland gorilla occupies the


Habitat

mountain region

Mountain gorillas

from 2 000

132

in

the Virungas occur at altitudes

m to 3 600 m, with

occasional excursions

of

Kahuzi-Biega. Here habitats

vary from dense primary forest intermixed with

bamboo,

to

mesophytic (moderately moisti wood-

Eastern gorilla [Gorilla beringei]

swamp and

Cyperus ICyperaceae)

land, to areas of

species of

Basella (Basellaceael,

Brillantaisia

Ipomea

peat bog, with alpine and subalpine grassland at

(Acanthaceael,

higher altitudes; patches of open vegetation also

(Convolvulaceae), Laportea (Urticaceae), Mimulopsis

occur at lower elevations.'"

(Acanthaceae),

Clitandra (Apocynaceael,

Mormodica

(Curcurbitaceaej,

Myr/anfhus (Moraceae), Palisota (Commelinaceae),


Triumfetta (Tiliaceae), and Urera (Urticaceae)."

Diet

Occasional items that mountain gorillas have

Mountain gorillas
Mountain gorillas are large-bodied herbivores;

in

the Virungas, they feed almost exclusively on the

leaves and stems of

harvested

the dense

in

supplementing

more
this.

herbaceous understory,

with bark and roots.' '"

this

Bwindi mountain gorillas

the

contrast,

Around the Karisoke Research Center

eating

gorillas

families.
of

in

in

the

species from

plant

18

linderi (celery, Umbelliferael; the

stems and roots

Laportea alatipes

of

and the stems and roots

Urticaceae);

Urtica

of

(stinging nettle, Urticaceael; as well as

massaica

leaves of

Carduus nyassanus

alatipes,

L.

and the leaves

Asteraceael,

(thistle,

(nettle,

ruwenzoriense (galium

which are speculated

cocoons

of unspecified origin];"-

""'

"^ at Karisoke

Research Center, subsoil sediments


times per year, possibly as
iron;'"

source

of

dung;'"'"'" and rotting wood."''"

been shown

to vary

and abundance

according

of food

the distribution

to

resources

that, in turn, vary

according to altitudinal and climatic factors.* For

groups

example,

at Bwindi,

altitudes

consumed more species

(UO versus
living at

62)

and

of gorillas living at

fruit (36

versus

higher altitudes. There

variation in the diet of the

is

mountain

111

is

high

seasonally and
gorillas

when

it

in

Rubus

spp.

of the diet of the

Bwindi gorilla varies

it

is

of

and

mountain

gorillas
of

at

Bwindi

than

mountain gorillas

at

A silverback male

eating

Myrianthus fruit.

Redmond

fluctuates

consumed

heavily by

mountain

abundant.'"^

A number

of feeding

is

number

similar between eastern lowland gorillas at Kahuzi-

Bwindi and the Virungas."

Its availability

seasonal

gorilla in parts

species eaten and the degree of frugivory are more

between the populations

and

shoots.'"''"

protein.

than those

little

available throughout the year,"" while the fruit

component

tan

Bamboo

lower

of fibrous food

the Virungas, probably because most of their food

Biega

bamboo

flexibility;

within both the Virungas and Bwindi their diet has

is

(berry, Rosaceaej;

(especially)

six

of

the leaves of G. ruwenzoriense, Arundinaria

P. linderi;

to

sodium or

Mountain gorillas show dietary

over the course of a year' The total

the stems of

five

Galium

gorillas have a preference for:

alpina (bamboo, Poaceael, and

to

Mountain

of

vine, Rubiaceae).

all of

have a nutritional function, include insects (ants and

of

study recorded mount-

38

In

"" '
These included the stems and roots

Peucedanum

the

live

and take advantage

habitat,

fruit-rich

Volcanoes National Park,


ain

and shrubs

herbs, vines,

been seen eating,

techniques have been observed; these are interpreted as measures to avoid injury from leaves

bearing stings or sharp hooks."' " As a result of the

low quality and poor

digestibility of

mountain gorillas

diet,

in

least half of their daylight


of the

much

of their

the Virungas spend at

hours feeding, and

much

remainder resting.'"

The

gorillas of the Virungas

and

of

Bwindi both

require abundant quantities of easily harvestable


plant material.'"'

The

gorilla

habitat around the

Karisoke Research Center contains


fruit,

'^

Bwindi
diet.'"'

as

is

reflected

fruit is
'^''

consumed

'^'

in gorilla

little

edible

diets there, while in

an important component of

gorilla

The most important fibrous foods

by the Bwindi gorillas also differ from

those consumed

in

the Virungas,

and include

133

World Atlas

of Great Apes and their Conservation

Eastern lowland gorillas

The varied

diet

lowland gorilla

eastern

the

of

includes a wide range of plants, their

and bark as well as

leaves, stems,

and other insects. Seasonality


use

fruit,

diet

in

and habitat

greater for eastern lowland gorillas

is

than

forests

altitude

mountain

for

Eastern lowland gorillas eat more

mountain

Bwindi

but

gorillas,

low-

in

gorillas.'"'

than do

fruit

not

When

western gorillas."'

seeds,

ants, termites,

much as

as

fruits are scarce,

eastern lowland gorillas travel less and increase

consumption

their

Large quantities
eral types of

fruit,

herbaceous vegetation.'"'

of

bamboo

of

shoots, as well as sev-

are eaten seasonally by eastern

lowland gorillas of the upper altitudinal reaches of

Kahuzi-Biega."

'

These

gorillas also occasionally

teed on ants, but have not been observed eating


insects as often as have eastern gorillas

more than

forests. Insects are never

in

lowland

minor part

of

the diet for any gorillas.' The ant-feeding sites


Martha M, Robbtns

A gorilla

infant,

Mountain gorillas appear

Bwindi

to

visit

feeding

recent use and those

Impenetrable National

areas that have received

Park, Uganda.

that produce nutritious food.' Foraging areas with


less

little

abundant high-quality

where renewal

food, or

rates are lowest, are visited less frequently than

When bamboo

other areas.
large

bamboo. As bamboo declines

on

exclusively

shoots are available

are sometimes eaten

activity
in

in

and expand the foraging area

In

Nutrient supply does

not appear to

mountain gorillas

in

be a

the Volcanoes

National Park."" While food abundance varies over


the range, no areas are so productive that

advantageous for

gorilla

groups

to

it

would be

ranges may overlap by up

resources

of

within groups
of

is

to

some

especially

in

ority

may also

but

is

100 percent,' ' with

that overall feeding competition

also rather low, and so the costs


'"

competitive disputes do occur,

of

in

The

of fruit

the environment,'"

be influenced by social factors such as

in

complex and changing


diverse ranging behavior,

groups generally spending more time


areas. '" Solitary males

have larger

home ranges

published

data

for

in

food-

the Virungas

in

than would be expected

for a single individual;"^'" there are

no equivalent

Food

Bwindi gorillas.

important Influence on the

movement

is

an

patterns of

males, and other gorillas are not always


"='"''

lone

avoided.'"'

Eastern lowland gorilla groups


forest have

to food, and there are weak dominance


among females.^' '"'" ' Hence, group
may carry some costs for lower-status black-

the size of

home ranges of 13-17


their home range in

forest

living

average day journeys

IS

lowland forests.

in

in

km'.'"'

montane
Although

lowland tropical

unknown, they are known

hierarchies

backs, females, and juveniles.'"'

year.'"

mainly deter-

Is

and abundance

distribution

silverbacks."^ These

larger groups.'" Silverbacks have pri-

access

In a

of a

"^"""^

about 5.5-11.1 km'.

may use 20-40 km'

factors are reflected

rich

social foraging are also likely to be low,'"

although

group

competition for mates or the mate-guarding tactics

Vi/ith

abundant, evenly distributed food

means

mined by the

home

the groups tending to avoid one another."'^ The


availability

gorilla

ranging behavior of gorilla groups

establish and

defend exclusive foraging zones."' Instead,

home range

the Virungas, the typical annual

and herbaceous vegetation

covered each day.'"'"

limiting factor for

trails

Ranging behavior

Bwindi gorillas

preferred foods are scarce, the

gorillas alter their diet

trees. Signs of feeding

valleys and swamps.'"'

bamboo areas and consume other herbaceous


all

in

have often been observed along gorilla

mountain

When

primary or ancient secondary

in

eaten on the ground, although leaves, bark, and


fruit

abundance, the gorillas move away from the

foods.

136

been found

all

in

mountain gorillas teed almost

quantities,

have

forests on ridges or slopes. Most plant parts are

montane

to

have shorter

forest than in

Eastern gorilla [Gorilla beringei]

occur

Ecological role

Not only

systems
to

behavior adapted

gorilla

Is

which they

In

to

the eco-

large, heavy,

dexterous animals that consume a

and

of foliage,

lot

they also change the structure of vegetation by

stem densities

foods increase
It

is

some herbaceous

of

the aftermath of gorilla feeding.'

in

whether there

not certain

Is

a positive-feedback

mechanism through which gorilla


more

regrowth and

This can stimulate

it.

productivity;

in

activity leads to a

edible plant community'"^'"

More than 30 years

made mountain

gorillas

social

patterns, and

the

AA and

species" Isee also Boxes

been reported

gorillas have

and the consumption

to

of fruit by

5.11.

Western

disperse seeds,'

eastern lowland and

may

Bwindi mountain gorillas suggests that they

Eastern gorillas share their habitat with


other large

mammalian

herbivores, and so might

them

be expected to compete with

Mammalian herbivores

for

food.

the VIrungas include

In

buffalo {Syncerus caffer] and

phus

bushbuck [Tragela-

scriptus], but these are not thought to have

Impact on the mountain gorilla

significant

'"'

population.""'

"

Other herbivores, such as

black-fronted duiker [Cephalophus nigrifrons] and

Information

the
to

Given

is

in

the

from

available

other gorilla populations.

eastern gorillas

multimale' group),

may

contain only

or

may

(in

consist of males

only.'""^ Most comprise a single dominant adult

male or silverback,

three or four

typically with

females and four or

five offspring."'
in

'"'

Over the

the VIrungas, between 10

percent and 50 percent of mountain gorilla groups

have been multimale," while at Bwindi about 50

percent

of

groups

are

multimale."

About

10 percent of eastern lowland gorilla groups are

multimale.

If

gorilla dies In a

the

dominant male mountain

one-male group, the group may


happen

A male

silverback

multimale

eastern lowland gorilla,

group, however, one of the subordinate males can

Democratic Republic of

disintegrate; should this

in

take over leadership and the group


Elizabeth A.

may

then stay

the Congo.

WiUiam5on

Elephants have considerable potential

gorilla.'"'

to

178. 183

gorillas

mountain

dietary overlap with the

little

UO.

between

one mature male, several mature males

the African forest elephant [Loxodonta cyctotis],

show

of

118. 134,

their

history

life

one important question

past three decades

also play this role.

ecology,

variability

Karisoke applies

best-studied

known about

demography 115.

ecological

Groups

of the

Is

feeding

behavior,

extent to which

regeneration and on the diversity of tree

one

primate species.''^ Much

both

are likely to have an Important impact on patterns

research at the Karisoke

of

Research Center established by DIan Fossey has

different habitats,

many forest communities, primates act

of forest

also have been

Social behavior

as seed predators and as seed dispersers; they

In

may

the VIrungas, and

from Bwindi.

but gorillas also help

live,

shape these ecosystems. As

trampling

lost

Impact the food supply

of the

mountain

gorilla,

but their

numbers are so low as

effect.'"'

Mountain and eastern lowland gorillas

to

have

little

real

are sympatric (occur together! with chimpanzees


in

some

overlap.

and their diets are known

areas,

'''"''

to

Although one competitive encounter

between chimpanzees and the Bwindi mountain


gorillas

has been observed, different foraging

strategies are employed

there

is

little

between them.'^'
patry with

It

has been suggested that sym-

chimpanzees may have promoted

eating strategy

niche

by these species and

evidence of feeding competition

in

away from

gorillas,

a leaf-

moving their feeding

that occupied by chimpanzees^"'

(see Box 8.1).

The only known predators

humans

and

leopards

of

[Panthera

gorillas

are

pardus]"''

Evidence of attacks by leopards on western gorillas


is

outlined

in

Chapter

7,

but these cats no longer

135

World Atlas of Great Apes and

their Conservation

intact."^ This pattern

seen

in

western

groups are extremely

Group size

is

"" "'

to that

among which multimale


eastern gorillas;

53 individuals have been

general, median group size

In

is

eastern and western gorillas,

similar for both

females with them from the natal group.

and transferring

to a

new

group), and secondary

dispersal (subsequent transfer to yet another


group),

occur among female eastern gorillas.

Females have also been known

remain and

to

61, 13i, 169. 183

re-

across various habitat types and the different diets


associated with them.' "" -' In the Virungas,

produce within their natal group

median and mean group

from their natal group alone, while female eastern

are eight and

size

11

individuals respectively (see Table 8.21. At Bwindi,


a
In

mean group

size of about 10

has been reported."

the area surrounding Tshivanga

the

mean group

solitary

group size

in

males)

only three

almost 10." Mean

is

the highland sector of Kahuzi-Biega

in 1996.^'

1990, to 10

group size

Kahuzi-Biega,

in

size of eastern lowland gorillas

(excluding

decreased from about

6 per group in

978, to

Other studies indicate a

seven animals

of

in

Kahuzi-Biega

approximately

is

Upon

1:1.

reaching maturity, most males and females leave


the group

which they were born

in

(their

natal

can attract females and establish their

own groups;

males form all-male

occasionally,

Virungas usually transfer

lowland gorillas sometimes transfer with another

female and their offspring."'

a female

If

pregnant

is

when she transfers to a new group,


that the new silverback will kill the

or has an infant

there

is a

risk

infant.'" Infanticide
in

has been observed occasionally

eastern gorillas,'""'^ although not

stance of transfer with an

infant'-'"

Female transfer could

(see

offer

every

in

Box

in-

8.2).

number

of

possible advantages, such as the opportunity of

into a

small or new group; avoidance of inbreeding;

increased choice

of

mates; improved reproductive

success; reduced feeding competition; or improved


protection against infanticide."'

Females may have preferences with regard

group). Males that emigrate usually remain solitary


until they

of the

Female

higher social rank,"^ especially following migration

Virungas and

ratio at birth in both the

mountain gorillas

mean

Kahuzi-Biega but

in

the adjacent Kasese region."

in

The sex

to

mates, and this choice

male

behavior.'""

may be

From the male

influenced by

point of view, good

groups. After emigration from the natal group,

relationships with females are important to mating

some males spend

a large proportion of their time

access and breeding success, as a female

alone, although

the

group.^"'^'

migrate

may

It

into

is

in

home range

of their natal

very unusual for fully adult males to

other groups.

53,

118, 195

115,

Young males

to

Its

leadership,'" "^ '"

groups, but not

all,

may

maturing and remaining

in

Most multimale

be the result of males


their natal groups,"^

and

essentially controlled harems,


fore afford to

Both aggressive and

oppressive ones.

affiliative

Males have been seen

to direct

aggressive displays

toward females, and females to appease those

males (see Box

displays and their impact on female

young male remains

in

his natal

group or emigrates could be determined by a range


of factors including

changes

in

social relationships

and demographic structure, such as the availability of

death

gage

parent, or disintegration of the natal


'"^"''
group.
Males that develop strong affiliative
(friendly] relationships with the

to the leading

therefore

dominant silverback

more

likely to

be close

male during adolescence, and are

more

likely

to

remain

in

group. ^^ Male eastern lowland gorillas

their
in

natal

Kahuzi-

Biega rarely stay with their putative fathers but

in

although the reasons tor these

mate choice

nonaggressive behaviors toward females,

possibly to

maintain proximity with females.'"'

Females may sometimes intervene

in

an attempt

end aggressive interactions between adult

mating opportunities within the group, the

of a

while they are infants are

8.31,

remain unclear'"' Males may also vocalize and en-

often, but not always, the case.'""

interactions

between males and females have been observed.

adult males. Genetic studies confirm that this

Whether

tree

males cannot there-

make them unduly

are therefore believed to contain several related


is

is

leave the group. Although gorilla groups are

also stay within the natal group and eventually

inherit

136

taking

^'

Both natal dispersal (leaving the natal group

among

to

own groups, sometimes

instead form their

rare.''"

variable

groups ranging from two


observed.""'

marked contrast

is in

gorillas,

Silverback males
interact

much

in

mixed-sex groups do not

with each other but,

the behavior tends to be

aggressive than
of

to

males.'"''

affiliative,

when

they do,

more competitive and


presumably as a result

competition over access to mates.""

Affiliative

interactions are rarely seen,'" but occasional co-

operation by males within the

been observed, apparently

to

same group has

prevent females from

leaving the group.'"' Relations

between silverbacks

Eastern gorilla [Gorilla berincei]

Box 8.1 COEXISTENCE OF GORILLAS AND


CHIMPANZEES

anzees [Pan troglodytes schweinfurthiil tend


avoid nesting

to

fruits of the

those trees with ripe

in

type preferred by gorillas." Eastern lowland gorillas


Gorillas

and chimpanzees

forests

in

existence

many

live

together

in

the

same

parts of equatorial Africa, a co-

known as sympatry. As they are so

how do

to one another,

manage

they

similar

to coexist

tend

extend their day-journey length during the

to

fruiting

season

in

both lowland and montane forests,

while sympatric chimpanzees tend to stay

small area, continually


'"'

'"'''

Such differences

without one species displacing the other'' Earlier

trees.'-

studies"- '"

patterns, and nesting-site choice

behaviors reduced competition through

ranging
'niche

suggested that their different diets and

Fruit-eating

differentiation'.

chimpanzees tended

to

range

IfrugivorousI

primary forests and

in

stay on the dry ridges, while leaf-eating Ifolivorousl

secondary regenerating

gorillas tended to

range

forests and stay

the wet valleys. These ecological

differences

and

to

in

were thought

to affect their societies,

determine their densities

habitat.
of

in

The dynamic

in

chimpanzees was therefore thought

of gorillas

were associated with

More recent
that there

is

in

both diet and ranging. Western

and range

their diet,

in

gorillas

in-

primary forests

close proximity to chimpanzees.^^''^'

Western lowland

and

gorillas include fruits

consume

in

'"""' """""

gorillas

ranging

diet,

in

may

limit

compe-

and chimpanzees.

and chimpanzees occasionally encounter

same

each other

in

Biega and

Bwindi, with

the

Kahuzi-

fruiting trees at

most encounters being

tense but peaceful.''*'^"* At Ndoki.

in

Congo, typical

encounters between western lowland gorillas and

chimpanzees are even more peaceful.'"

Many aspects

among

of

foraging

behavior seen

and chimpanzees may vary with

gorillas

environmental conditions, and the true extent

unknown. This

variability is still

tant in predicting

their folivory.

however, have shown

studies,

and eastern lowland


in

be caused

actually extensive overlap of gorillas

and chimpanzees

sects

to

more cohesive groups

between sympatric

Gorillas

different types of

'fission-fusion' social structure

by their frugivory, while the

tition

in

revisiting particular fruiting

fundamental

to

of this

be impor-

how gorillas and chimpanzees will

change wrought by humans, which

react to habitat
IS

is likely

to

wise conservation planning.'"'-

"'

Continuing research on eastern lowland gorillas

and sympatric chimpanzees


Bwindi

at

Kahuzi-Biega and

scope

will help to clarify the

for improving

the survival of sympatric great ape populations.


Juichi

plant foods as

Yamagiwa

diverse as those eaten by sympatric chimpanzees.


Of the fruit
gorillas at

are also

species eaten

by western

lowland

Lope National Park, Gabon, 79 percent

consumed by chimpanzees

in

A female

gorilla

and

infant,

Kahuzi-Biega

National Park.
Jijichi

same

the

Yamagiwa

forest.'" All fruit species eaten by eastern lowland

gorillas at Kahuzi-Biega are also eaten by sympatric

chimpanzees.'"' However, analysis of fecal samples


at

Kahuzi-Biega and Bwindi shows that there are

marked differences between the two species

in

their

reliance on particular fruit species, such as Ficus

spp. iMoraceael,
sp.

and Drypetes

Syzygium

sp. iMyrtaceael, Bridetia

sp. (both Euphorbiaceael.''"''"^

The presence

of gorillas is

choice of nesting trees

the

secondary forests

at

and blackbacks tend

thought

to influence

by chimpanzees.

In

Kahuzi-Biega, eastern chimp-

to

be weak.'"-"" Blackbacks

are subordinate to silverbacks, and generally spend


a lot of

Young, unrelated males that form all-male

and perhaps

to

do so

to

to increase safety

develop social

be more

affiliative

than

in

all-male groups tend to

among males

in

mixed-sex

groups, as measured by the occurrence of playing,

time on the periphery of the group.'"

groups are thought

Relations between males

skills,

from predators.'"

'"

close proximity'"""

grooming, and time spent

in

Homosexual behavior has

also been observed."^

Aggression

is

more frequent

in

all-male groups, but

137

World Atlas

of Great Apes and their Conservation

Box 8.2 INFANTICIDE IN GORILLAS

cases

Few behaviors observed


have led

in

the animal kingdom

more heated debate concerning

to

its

function than infanticide (the killing of young from

same

the

Why would

species).

the

of

killing

The

risk

infanticide

of

played a large role

thought to have

is

shaping the social behavior

in

and group structures observed

Where there

species.'"

western

in

group disintegrations.'"

many primate

in

only one male per group,

is

dependent young evolve as an adaptive strategy'

females can exert mate choice by transferring

The

between social

is

prevailing view

that

is

by males

infanticide

related to competition over access to females,

with

line

in

Specifically,

sexual-selection

the

if

a male

hypothesis."

unweaned

kills

offspring of

because

units;

infanticide, the opportunity for a

without risk

is

limited to the brief time

she does not have a dependent

the

of

female

risk

of

to transfer

window when

offspring.'^'

multi-

other males, and thus shortens the time that he

male group structure

must wait

the event of the death of the leading silverback,

to

impregnate their mothers, he

own

increase his
that of other

infanticide results

in

in lost

will

compared

reproductive success

males who do not follow

Given the cost to females

to

this strategy

reproductive

effort,

a conflict between the sexes.

Infanticide occurs rarely,

but over the past four

decades has been suspected or obseA'ed

more

in

than 40 species of primates, including gorillas.

Known
times

at

mountain

gorillas

were recorded

13

Karisoke between 1967 and 1988; these

all

were indeed cases

of infanticide,

for at least

37 percent

infant mortality during this period.'"

these cases occurred

of

when

of

The majority

the

mothers

of

the infants were not accompanied by the group's


silverback, typically

because he had

died. This

sug-

gests that an important motivation for females to

form long-term associations with males

is

to obtain

protection against infanticide.


infanticide

Is

exclusively

Interestingly,

in

Kahuzi-Biega,

among

Karisoke have been almost

at

multimale,

no group disintegrations

males been observed or suspected."


rarely

gorillas'

female eastern

Infanticide

has

been observed during encounters between

groups, and male eviction and group takeovers by

ing the

the

new

time

of

females

'^'

have

been observed

not

simultaneous transfer

of several females,

silverback killed one unrelated infant at the

and (despite the

transfer,
in

efforts

of

the

the group to intervenel killed two other

which occurred

infants shortly after their births,

only a few
In

in

'" Recently, at
Kahuzi-Biega, follow-

months

after the transfers.

addition to

has implications
of a silverback

universal

take

male."^ '" Since the late 1980s, while the gorilla

groups studied

gorillas.""'

If

likely to

group disintegration and infanticide by an outsider

cases, and one unsuccessful attack inferred from

would have accounted

is

in

over the leadership of the group; this prevents

extragroup males

this

advantageous because,

another (often related! adult male

comprised three observed cases, nine inferred

wounds.'"

is

have occurred, and neither have any infanticides by

or probable infanticide and attempted

infanticide in

one individual

its

impact on

for population

represents
in a

sociality, infanticide

dynamics. The death

initially

population.

If.

the loss of only

however, he was

the leader of a one-male group, his death

is likely to

unweaned

offspring.

lowland gorillas with dependent young have been

lead to the deaths of

obsen/ed unaccompanied by silverbacks

This impacts overall infant mortality, future births,

for

many

months. Females have also transferred between


social units with
killed,

unweaned

at

disputes between males

in

were not

have been

all

his

group age structure, and the rate

in

mixed-sex groups (when

more serious and more

wounds.'" This difference

result of competition

is

likely to

probably a

between males over mating,

an issue that does not arise

in

all-male groups.

such as those

of the

mountain

offspring, social

bonds between females tend not

to

population

gorilla.

Martha M. Robbins

be well developed. Females


natal group, so

commonly

leave their

complex social networks between

females do not occur. The female coalitions that

do emerge, allowing

common

defense against

aggressors, are thought to be more

Apart from those between mothers and their

of

growth, which can be critical for small populations

Kahuzi-Biega."' On the other hand, two

they occur) are


result

infants that

but three cases of infanticide

observed

138

have been inferred

of infanticide

gorillas following

common among

related than unrelated individuals.'"'" Males fre-

quently intervene

in

conflicts

between females, thus

Eastern gorilla [Gorilla beringei]

female coalitions. Such

limiting the effectiveness of

interventions involve only moderate aggression, pose


little

risk to social relationships with females,

may help males to retain mates by maintaining


own status and control over the group.'"
Immature

receive defensive

often

gorillas

support from their

and
their

mothers,

but

from

rarely

unrelated adult females. Juveniles rarely receive

support

consistent

however,

if

from

even

mothers,

their

they behave aggressively toward larger

opponents.'" During infancy, gorillas often develop

an attraction

may

to the leading

buffer the

male

who

of the group,

young animals against aggression

from others, serve as a spatial focus

for

young

animals, and provide an attachment figure as the

maternal bonds weaken.'"" The behavior


toward infants and juveniles
no great effort
protect

is

immature

but provide

little

who behave
conflicts

is

male

of the

though

paternalistic,

put into this."' Adult males


gorillas against larger

support

immature

to

individuals

mostly

intervening

aggressively,

between immature peers only

may

opponents

to

in

maintain

Virungas

(A.

6 versus 3.9 years].

infant die, this interval

mother

conceive again within three

to

months. Infants are

control."'

typically

four years,"' but

Reproduction

Social

than one female lare polygynousl. Generalizing

change during an

from the Virunga

gorillas,

seems

it

that female

mountain gorillas reach sexual maturity around the


of six

the

first

and a

halt years (5.8-7.1 years).

Between

bout of estrus-like behavior and the

conception there
that

is

phase

two years. ^'

lasts

among young

and group composition

may

'"

of

adolescent

first

sterility

Although less regular

among

females, the menstrual cycle

rank

different relative position within the

expected

to

attractive to

males

around

at

mid-cycle, for one to four days." The gestation

period lasts about eight and a half months.

Mating or mating attempts occur

at

^^^'^' '"

times during

menstrual cycle and pregnancy when

estrogen concentrations are highest.""

Mountain gorillas do not have a

presumably because

'"

change

multimale groups
only

group

of

mountain

gorillas,

Virunga

National Park.

that individual's reproductive

exist.

male present does

when conception

more

all

is

is

'""

exceptionally rare

in

sometimes even

is

longer than that

of

mountain gorillas

slightly

of

the

mountain

gorillas.'"
try

to

Females sometimes mate with more than one


male,

occur, and

is

of

proximity to females at mid-cycle.'"

interval lasts approximately four years, as

amenorrhea). The recorded interbirth

in

multimale groups, males often

Harassment

interval for the eastern lowland gorilla

males

Mating with individuals from other

coercion.'"

suckling young

although

more mat-

subordinate males do sire a proportion

offspring.'^'

susceptible to respiratory infections."^ The inter-

still

likely to occur,

that

may be

gorillas are not fertile while

multimale

In

to participate in

period. This

the animals are colder and

the mating.

subadult females.'" Genetic studies reveal

remain

and

many

one-male groups, the

ings with adult females, and subordinate

groups

lApril

In

with

of the lack of seasonality in

months

group can be

groups, subordinate males do mate, including at

In

are highest during the wettest

Assuming

sidered to have a one-male mating system,

season,

birth

food availability. Infant mortality rates at Karisoke

(lactational

Part of the Mapuvra

Although mountain gorillas are con-

strategies.

dominant males tend

birth

variation

is

individual's lifetime.

times

when

at three or

both

most receptive and

May],

six

to

in

there

adults has a median length of 28 days; females are

both the

weaned

its

directions.

Successful gorilla males typically mate with more

age

Should an

shortened, allowing

is

in

the

same mid-cycle

voluntary or the result


of

of

male

copulating males can

often but not always practiced by

dominant males.'"
Eastern lowland gorillas share
ductive

characteristics with

many

mountain

including a sterile subadult period

in

repro-

gorillas,

females, the

139

World Atlas

of Great Apes and their Conservation

Table 8.1 Eastern lowland and mountain gorilla populations

Subspecies

Approximate

Approximate area

population size

Mountain

gorilla IVirungasI*

380"-"

Mountain

gorilla (Bwindi)

320""

Eastern lowland

occupancy Ikm^)"

of

375
215

?'

gorilla''

15 000

17 000 + 8 000"

See also Table

8.2.

b See also Table 8,3


c

No

data; fieldwork

d Estimate based on

was
1

being undertaken

in

2005

to

998 survey data, obtainec prior

estimate the extent of the decline,

to

outbreak of war

^'

the area

in

Table 8.2 Mountain gorilla populations of the Virungas (1971-2003)

Census

Total gorillas

Estimated

Number

Mean

counted

population

of social

group size

years

umber

Multimale

Immature

of solitary

groups

individuals

size

groups

males

(percent)

(percent)

1971-1973'^' "

261

27A

31

7.9

15

42.0

39.8

1976-1978""

252

268

28

8.8

39.0

35.8

1981^

2^2

254

28

8.5

40.0

39.7

1986"^

279

293

29

9.2

1!

8.0

48.2

,98912'

309

324

32

9.2

28.0

45.5

2000"

359

359-395

32

10.9

10

52.9

44.7

2003"

380

Adapted from Katpers Je(3M2003l.

Table 8.3 Eastern lowland gorilla populations

Geographic region

Estimated population size

Estimated population size

(2001-2004)

Kahuzi-Biega National Park and

(1994-2000)

15703

17 655-224911
(1994-19951"'"

present 120051

adjoining Kasese region

Tayna and other proposed community

,a

050 1700-1 4001 120041"

resen/es

Maiko National Park

assumed present

llombwe Forest
Northern bank
(north of

859 1462-1 1351(19961"

120051

present 120051
of

Lowa

155 1516-1 7961119991''

13 10-261(1998)"

20 120041"

present'"

2810-33111988-19981"

small"

River

Kasese region]

Mount Tshiabenmu, Virunga National


Park
Masisi Imcluding Shingisha Mabeshil

Mbohe, North Kivu

'' indicates that no data are available

Adapted from

uo

Hall. J

S, e(

a( 119981

and

later sources, as cited

the table

Eastern gorilla [Gorilla beringei]

age

at

parturition

first

interval,

and

(giving

interbirtti

birtln),

infant mortality rates/"'

DRC

parts of the Virunga Massif, seven habituated

groups had declined from a

gorilla

total

number

of

103 individuals to 66 between 1995 and 1998, but

showed an

Nest building

weaned immature

Adults and

each

night,

gorillas build nests

which they sleep. Unweaned

in

off-

The

gorilla defecates either in

or next to the nest, and the size of the dung

The increased numbers

is

mountain gorillas

of

revealed by these censuses should be viewed with

some

caution because nearly

all of

the population

the

Research/Susa

''""

age

directly proportional to the

growth can be attributed

of the gorilla."

Counting and measuring nests and dung can


therefore provide information on the

group and the age class

gorillas in a

vidual using each nest, so

census method.

In

it

is

number
of

of

the indi-

commonly used

the Virungas, mountain gorillas

almost always make nests on the ground, while


about half
in

the lowland tropical forest of Kahuzi-Biega are


in

trees.

montane

the

In

made on

Kahuzi-Biega, most nests are


but,

forest of

the ground

even here, immature gorillas tend

nests

trees

in

more

if

thought

the

make

gorillas tend to nest in

groups sllverback has

be a result

to

to

frequently than do adults;

more immature and female


trees

to

section of the Volcanoes National Park, an area that


is

relatively well protected,

and which

believed to

is

be a particularly good gorilla habitat. Other sectors

known
number of

are

work

to

have experienced a decline

gorillas," so there

is

still

the

in

conservation

to do.

nests of eastern lowland gorillas

of the

constructed

of the

Virunga gorillas recorded 380 animals."

spring share the nests of their mothers; otherwise,


gorillas sleep alone.

between

overall increase from 66 to 86

1998 and 2002." The most recent census

died. This

is

Mountain gorillas

in

Bwindi

The small Bwindi mountain


appears

gorilla population also

A survey

be stable.

to

the early 1990s

in

found about 300 animals, '= " which was confirmed


by a complete census of the entire park

199Ds," and raised


in

to

the late

in

about 320 by another census

2002."

of their vulnerability to large

Eastern lowland gorillas

terrestrial predators.'"

The

area known to be occupied by eastern

total

POPULATION

lowland gorillas declined from about 21 000 km^

Status and trends

1963

The population

of

mountain gorillas

of the

Virungas

to 15

geographic range, calculated by Butynski from

has been monitored since the 1970s. Fewer data

historical

are available on the status and trends of mountain

illustrates the

gorillas at Bwindi, or of eastern lowland gorillas.

Recent estimates

of overall

numbers

of

eastern

to

86 percent

Mountain gorillas

The mountain

in

gorillas of the Virungas have

A summary

population estimates can be seen

These data show

into the early 1980s, with


in

the

mated

DRC
to

of

section.- ""

in

1973," and 254


in

in

population estimate

in

1981.=

in

esti-

the late

The 1989

the Virungas counted

309 animals and estimated 324

of

and

most reduction occurring

mountain gorillas

observation

Table 8.2.

The population was

contain about 450 gorillas

1950s,'" 275

census

in

a decline through the 1970s

to

be present.'"

2000, based on repeated

17 habituated groups and

infor-

of

This

fragmentation of popu-

By the mid-1990s, there were

in at

eastern

8 000)

least 11 subpopulations, with

Kahuzi-Biega and the adjacent

DRC."'"

More recent events

been

of selected

of

be about 17 000

living in

Kasese region

the Virungas

studied for over AO years.

degree

lations at that time.

estimated

was 112 000 kml"

data,

locality

lowland gorillas

gorillas are given in Table 8.1.

in

000 km^ by the early 199Ds. The overall

in

Kahuzi-Biega and the

surrounding region, however, indicate that the


species has undergone a substantial decline

numbers'""^
gorilla
is

(see Table 8.31. Access to

range has been

difficult in

much

The

available

consensus

information

is

among

workers that a drastic decline

very limited, but there

is

population has occurred. This

is

combined

demand

ore

effects of the rise

(discussed

in

more

in

in

the

recent years, and

only just becoming possible again.

field

of

in total

attributed to the

detail

warfare that engulfed the whole

for 'coltan'

below)
of

and the

the eastern

mation on 15 unhabituated groups, suggested that

lowland gorilla range from the late 199Ds onwards;

the Virunga population of the mountain gorilla had

armies, rebels, refugees, and miners

further increased to between 359 and 395."

land and

In

the

all lived off

the

consumed bushmeat."'

141

World Atlas

of Great Apes and their Conservation

Box

THE VOCAL BEHAVIOR OF MOUNTAIN

8.3

GORILLAS

study groups at Kansoke. adult gorillas vocalized

about once every eight minutes. Over half

occurred as part of an exchange,

calls

Mountain gorillas use a variety

of vocalizations to

individual.

group- Calls aimed outside the group are given

gorillas usually give

males

primarily by adult

in

response

to potential

such as a human hunter or a


These

silverback.

more

intimidating

and screams; they are sometimes accom-

roars,

When

panied by a charge.

encountering another

group or a lone male, adult males also give a form


of

long

vtfhich is a

call',

combined with displays such as

usually

hoots,

series of loud, resonant

'"

chestbealing or ground thumping."-

more frequent and

energetic, but far


of

these signals occur

of the

calls

is

human

an

infant that

has

to

age and dominance

frequently than do adult females,

'close calls' that gorillas give throughout the day in

common

various nonspecific contexts. The most

of

these signals are atonal, belch-like grunts, usually of

one or two

syllables, that

male clearing

his throat.

calls, similar to
If

case, the vocalizations

the time, however,

Biega;

in

location

in

population

in

is

singing."'"

viewed as a form

calls'

are the

lack of

two

decline

lowland gorillas
the

in

same

the lowland sector of Kahuzi-Biega

mammals
meat

of this

most
In

The eastern lowland gorilla


wardens

even greater casualties; a crash


of large

of

is

to

have suffered

in all

populations

inferred from the reported

these formerly abundant species

bushmeat supplies

what purpose the signal

The syllabled grunts are

particularly

The animals grunt most frequently

enigmatic.

during feeding, while traveling, or resting. Calls

evoke either no discernible response

or. at

most, a

vocal answer from another animal."'"

While acoustical analyses indicate that

down from 245

believed by the park

of

grunts are individually distinctive (suggesting that

the mountain sector of Kahuzi-

1999,

Most

what prompts

human

then 'close

996."'' "*

subordinance
'" '"

include

here only 130 eastern

remained

to vocalize, or

ser^'e.

to signify

signals."'

not clear

is

they are

The best-documented example


in

it

when

is

just displayed. In this

like

frequent social interaction between gorillas.

is

seem

and act as appeasement

One

which adult

calls'

vocal communication

population

intensely

in

Other 'close

human humming and

of social behavior,

turn

sound much

grumbles, and higher-pitched tonal

syllable-free

in

do more humming and singing.

obvious, but not exclusive, context

might

frequent, quiet,

are

mainly of syllabled grunts, whereas younger

gorillas

an animal

more mysterious are the

who

immature animals. The adult vocal repertoire con-

hum

accompany copulation."

who

males,

more vocal than younger and more subordinate

'chuckles' given only during play, or the staccato


that

status. Adult

dominate other group members, vocalize more

who has

mother, the breathy

that

other

correlate with

gorillas

of

near an adult male

whimper-

is

when

calls

the nature and frequency of 'close calls' are related

whimpers
Far

many

gorillas can recognize

each other from their grunts],

few features

sounds

As

far

as

we

of the

can

tell,

relate to behavior'"

grunts given during feeding

are the same, acoustically, as those given during


resting.

is

It

possible that these signals convey a

mostly ate large


relied

mammals; toward

upon small mammals,

conflict situation

birds,

has prevented

the Wildlife Conservation Society


a gorilla

offer a
In

survey

more

in

2004-2005;

solid estimate of

it

the end, they

and

of

The

surveys, but

was

coordinating

is

hoped

this will

remaining numbers.

summary, about 700 mountain

thousands

turtles.

field

eastern lowland gorillas

gorillas
still

and

survive.

Both subspecies have declined significantly

numbers. This process

is

in

ongoing (perhaps catas-

sold by hunters to coltan

trophically sol for eastern lowland gorillas, while

miners."'' At the beginning of the coltan rush, the

the mountain gorillas have been slowly increasing

miners

since the early 1980s (Table

in

U2

The vocal habits

females grumble or

often quite clear, as for

lost its

and exchange

and often

the nnildly aggressive 'cough-grunt', the


ing of

Some

key feature of this vocal behavior

observers, the

specific contexts

in

evoke specific responses. To

meaning

varied.

by a call from another

other aspects of their social behavior For example,

sists

Intragroup vocalizations are quieter and less

which

individuals are nearby, within 2-5 m.'

rival

convey alarm/threat and

calls

include various types of 'barks',

is

communicate, both within and beyond their social

danger,

in

was 'answered'

vocalization

these

of

in

in

the

lowland sector

of

Kahuzi-Biega

8.2).

Both the Virunga

Eastern gorilla [Gorilla beringei]

general message such as


or simply,

activity",

communication

am

"I

"I

am

here".

about

change

to

The function

of this

then depend on the context. For

will

example, during feeding


might be important

in

periods, vocalizations

the avoidance of feeding

competition.^'

seem

situations,

close

ordinating

group movement and

calls'

and

interindividual spacing

other

In

to play a role in co-

Toward

activity

the end of a midday rest period, resting gorillas

increase the frequency of their grunting, as


indicate that they are ready to

seem

be signaling their

to

end the

move

'intent' to

to

if

They

siesta.

on, but

wait to do so until they have heard from the rest of

the group. Even


but lying

still,

rest period

is

when

the animals are doing nothing

an observer can often


about

when

tell

the

end, just from the increase

to

in

'conversation'.'"

our data on vocal communication

All

comes from

wild

mountain

the

Virunga Volcanoes. Studies

of

gorillas

suggest that the vocal repertoires

the

We

in

the wild

in

of other

lations of gorillas are generally similar

much more

the

western gorillas

and preliminary observations

captivity

in

of

popuhave

still

however, about gorilla vocal

to learn,

communication.
Kelly

Above:

A young

J.

Stewart

sitverback hooting during a chest-

beating display. Betow:

An

adult female and

silverback playing; they have just sat back from

some

gentle wrestling. The female

chest.

They both have the open-mouthed

that

is

beating her
'play face'

accompanies the breathy pants known as play

chuckles. These vocalizations are characteristic

and very

specific to, play.

of,

They are given by young

infants upwards.

and Bwindi populations


classified

(separately,

of

mountain

because

gorillas

of the

were

uncertainty

The hunting

of gorillas for sale

heads, skulls,

feet,

as trophies (skins,

and hands -

sold, for

example,

over their taxonomic status! by lUCN as Critically

as ash-trays) emerged

Endangered, on the basis of their small population

continued until quite recently.'"' '" Occasionally

sizes, with

fewer than 250 adults

in

the

mid-1970s, and

each case;

individual gorillas that raid the crops of local people

as

are killed." For example, a young mountain gorilla

1998

estimate of population numbers."

was stoned to death in January 2003 when his group


damaged fields near the border of the Virunga

Threats

this habituated group,

Hunting

crossfire."

eastern

lowland

gorillas

were

in

classified

Endangered, albeit on the basis

of

the

National Park;" Rugendo, the previous silverback of

Gorillas are hunted for their meat, as


(particularly infants] for collections,

specimens

and as trophies.

had been

Infant gorillas have

killed in

been captured

2001

in

for sate,

or attempted sale, to public or private collections,

U3

World Atlas

of Great Apes and their Conservation

vsar^'iffv"

7=3rtirT

:';:i(K'

where

locations

is

it

abundant, including rivers

in

Kahuzl-Blega.'""" Professional hunters joined the

miners

to provide

lowland

gorillas

affected.'"'

"''

meat

them, and the eastern

for

Kahuzl-Blega were severely

of

More information on the decline

eastern lowland gorillas can be found


country profile

16.

were

Traditionally, gorillas

HOSTS. WHO
\TRrni\r:irMrYKiLi;;'n

\U-A

Chapter

in

of

DRC

the

in

rarely eaten In the

iiiLiK

eastern Congo Basin, which has given the eastern

amount

gorilla a certain

are weakest

ditions

of protection.

These

tra-

areas inhabited by the

In

eastern lowland gorilla and, as seen

Kahuzi-

In

Biega, are fast becoming a thing of the past. They

JOA-NNtCOT
MARTIN

were and remain strongest, however, around the

^^

Virungas and Bwindi, providing continued protection

SNHOBVAaD.8.on
A plaque

in

to the

War and political unrest

Bwindi

Wars

National Park

commemorates seven
those

hands

who

-I'SS-SS?"

of

and death can

gorillas as well as people,

kill

disrupt gorilla groups as effectively as

may

communities. Gorilla groups

died at the

of militia in

mountain gorillas there."'

March

response

1999.

losses,

to

sllverback,

it

does human

disintegrate

which can result

in

In

dominant

particularly of the

additional mortality

Gordon Milter/IRF

and declining populations. Armed


and many adults have been

while trying to

killed

protect their infants from this fate."'

mountain

infant

gorillas

serious problem

in

in

The capture

of

was

the Virungas

the 1970s, although

declined

it

greatly through the 1980s and into the 1990s.

were

1995, however, four adult gorillas

Bwindi,^ and

have been reports

there

killed

In

on both the

wildlife of

DRC, Rwanda, and Uganda.

The early 1990s saw the outbreak


in

of fighting

Rwanda, including within the Virungas; by

DRC and

Poaching leading

stream

of

and

surrounds. About half of Rwanda's

nine
for
in

in

the deaths of at least seven

2002

in

the Virungas;

Rwandan poachers were

two

to four

fined

that

late

killing

had been protecting

remains a threat
In

2003,

years each for stealing a baby gorilla

Volcanoes National Park, and for

gorillas

in

and imprisoned

response

In

two adult

it.'

Hunting

to the situation in

DRC from

the

1990s onwards, the United Nations Security

exploitation of natural resources In

cluded that the various armies active

DRC.
in

natural

five

was displaced during

of the

It

con-

DRC came
in

gorillas

and

In

the vicinity

In

Kahuzl-Blega." Soon

Rwandan refugees

DRC;

put

massive

habitats through

pressure

harvesting of

wood

cobalt,
of

the manufacture of mobile telephones

of

were located

Park buffer zone;


ted by

wood

conflict In

for fuel, increased hunting,

the four refugee


In

on

uncontrolled

disruption of migration patterns. During the

an alluvial ore

Into

fighting broke out

resources

niobium Icolumbium] and tantalum, metals that


are used

in

DRC were

Kivu

is

war

can

their

weapons; these were diamonds, copper,


coltan.""' '" Coltan

civilian

1998.

Refugees

Rwanda, three

and

DRC near

the 1996

either to finance themselves or to exchange for

gold,

Virunga National Park, and a further 332 000

having fled into

again

in

this conflict, with

860 000 refugees being concentrated

Council established an expert panel on the illegal

systematically exploiting

its

population

April

resulted

refugees pouring Into the gorilla habitat

after the 199(1-1995 influx of

the Virungas.

wars

populations, with a series of conflicts and

which have affected the people, landscapes, and

199i, this had expanded Into

to

and

conflict

toll

gorillas being taken for sale to private collectors.'^'

gorillas occurred

U6

unrest have taken their

eastern lowland gorilla and on the mountain gorilla

in

infant

of

political

camps

In

and

war

In

North

or near to the Virunga National

much

of

the park has been affec-

harvesting or poaching.'^' Subsequent

DRC

led to looting

and destruction

of the

and computer equipment. The ore has a ready

park's infrastructure, and the deaths of about 5

market, and

percent of the mountain gorilla population

Its

high value has attracted miners to

in

the

Eastern gorilla [Gorilla beringei]

These factors

Virungas.

led

Virunga

the

to

National Park being placed on the 'World Heritage


in

Danger'

list in

In

99A.'" As described above, hunt-

meat

ing for gorilla

as a result

Kahuzi-Biega has increased

in

war and displacement.'"'"

of

home

places and

have served as hiding

to gorillas

disturbance and hunting. This

nomenon

leading

retreats for rebel forces,

times

at

is

common

to

phe-

of Vi(ar in forests that straddle

international borders.'"
of the recent

Central Africa are unclear, and the

civil

and, despite intense military activity

in

in

1989

in

2000."

DRC, where most

The lowland protected areas

eastern lowland gorillas

of the

occurred during the 1990s, remain inaccessible


researchers so

it

The population

is difficult

in

of

to

to

assess their status,'"

the area around Tshivanga

In

all,

92 Congolese

1996 and 200^." During the conflicts

in

several workers from

their

Karisoke

lost

others were imprisoned, and the center

Rwanda,
lives,

itself

was

destroyed;"''"'" much more international atten-

was drawn

by the killing of eight tourists and

March

1990s, there appeared to be at least 57 gorillas

remaining

""'"'"

in

the early

in

wildlife.^'

wars

somewhat

numbered about 57

its

four guides at Bwindi by

hopeful sign relates to the mountain gorillas


the eastern Virungas. This small and

or the only

park staff are reported to have been killed between

in

it

first

while they were attempting to protect the area

wars

difficult to assess
Rwanda and DRC have made
how the mountain gorillas have fared,'" although
some censuses have been carried out." One

reestablish the park

to

" These were not the

park-service employees to be kidnapped or killed

tion

The long-term impacts

isolated subpopulation

limits. ^^'

and

addition to the influx of refugees, the forests

that are

Kahuzi-Biega boundaries

1999.'

mitment

of

Interahamwe

militia

in

Without the determination and com-

park rangers,

it

would be impossible

imagine the long-term survival

to

of the eastern gorilla.

Habitat loss or modification

The mountain and eastern lowland


surrounded by some
populations

in Africa,

of the

gorillas

densest rural

live

human

with up to 300-600 people per

square kilometer, and a correspondingly high

demand

for land

and

food."'^ '"

As a

result, gorillas

are increasingly confined to smaller and


lated

forest

increase."

more

iso-

fragments as human populations


Cattle herding in Uganda.

in

Gordon Miller/IRF

Kahuzi-Biega was relatively stable between 1990

and 1996"

but,

since then, two rebellions have

occurred, with large

numbers

gorillas being killed."^ "'

of

highland sector of Kahuzi-Biega

95 percent

of its

percent of

its

eastern lowland

Over just four years, the

more than

lost

elephant population and about 50

resentment

gorilla population. Local

toward the park and

its

authorities

may have

contributed to this illegal exploitation of wildlife

resources.'
Conflict can also deter international conser-

vation

organizations,

ments from

agencies,

aid

investing

and govern-

affected areas, leading to

in

frozen budgets, withdrawal of

reduction

staff,

antipoaching efforts, and the closure

Nevertheless,

some

organizations continued to

support park authorities

in

the Virungas throughout

the war, even though research

programs were

interrupted. Protection of the gorillas

has proved extremely


in

recent years, and

difficult

many

take tremendous risks

sometimes with

fatal

in

of projects.

in

in

many areas

and often hazardous

national conservationists

the course of their work,

consequences. Ten

staff

and

assistants of ICON, for example, were murdered,

apparently by militiamen

who had been

DRC

Rwanda, while surveying

since the genocide

in

hiding

in

U5

World Atlas

of Great Apes and their Conservation

Habitat loss, specifically forest clearance for

was one of the mam causes of population decline among mountain gorillas during
the 1970s." "* In 1968, more than one third of the
Rwandan Volcanoes National Park was excised for
agriculture,

an agricultural project."'

remains

Rwanda, and

in

Little

virtually

now

forest cover

no forest habitable

by gorillas remains outside protected areas." The

boundaries

protected areas are generally res-

of

pected by planners and farmers, so there has been


very

little

further habitat loss

presence, social

genocide, and war has occurred.'"

instability,

forest has also

and

for building

and

Rwanda, although

human

disturbance from increased

The

in

been used as a source

fuel,

and

accessed both for water

is

Mgahinga

agricultural and

Uganda,

pastoral activities and

hunting

were major pressures; incursions by


their livestock

has

used

common. The park

be

to

complex history

local people

designation changes,

of

game

having originally been defined both as a

reserve and a forest reserve, established

and 1939 respectively. The boundaries

in

425 m, on the lower slopes

what

now Mgahinga

is

of the

m, thereby

and removing some important

Mgahinga

After the

designated

in

was lowered

the

to

significantly increasing

that

game

was already

280

raised to

was

now attempting
In

to

it

the

high-altitude

forests of the

vegetation, and so might be able to coexist with


logging,

if

it

were not

for the associated hunting.'"

The Bwindi population

mountain gorillas

of

relatively well protected. Prior to the

of

and head-load extraction

980s,

wood, usually

of

branches, that can be carried on ones head!

was then

a forest reserve.

made

techniques

These nonmechanical

for very selective

and environ-

mentally benign logging. Nevertheless, only about

human

in

disturbance.'^'

Bwindi

No

is

entirely free of

data are available on

the intensity and distribution of habitat disturbance

since Bwindi was declared a national park, since


when antipoaching and other enforcement efforts
are thought to have led to much reduced levels of

disturbance."

Eastern lowland gorillas and their habitats


face similar problems of habitat loss, which add
to

the impacts of hunting that have been noted

human

population and the


is

serious and

ongoing pressure." The boundaries

game

an important area

of gorilla habitat."

com-

suggested that the rate of loss

of extractive

use

of

park

to the
is

balance the needs of the people

of

timber was permitted throughout the area, which

in

large

is

manual

the removal

(i.e.

no more than the quantity

Biega were altered

that a

its

Virungas. Gorillas often favor areas of secondary

corresponding need for land

1964,

into

unlikely that large-scale logging

is

in

contour

in

move

quickly

impact eastern lowland

above. The increasing

A community-based conservation program

and the

gorillas, but

would occur

'"

area were

resources occurs both within and adjacent


park.

will

commercial

that

The designated national

means

munity with a tradition

companies

forests.'" This could well

000

As DRC

rebuilt.

is likely

it

reserve boundary

park encompasses part of this additional


reserve area,"' which

area

its

area and including land

its

settled.

in this

stable,

10 percent of the forest

gorilla habitats.'

1991, people living

Meanwhile, the

evicted.

becomes more

past

National Park

Gorilla

boundary wall was

drystone

and the parks

killing two,'

1930

significantly reducing

removed the

soldiers

each

Gorilla National Park. In

was

Rwandan

in

three volcanoes

1951, the forest reserve boundary


2 730

park." After international

loggers and farmers,

logging

the Mikeno sector, also on

in

the

of

for

reserve were defined by the contour line running at


2

side

protest,

felling

Gorilla National Parl<,

May 2004

in

DRC

wood

graze cattle."'

to

In

and

of

"'

farmers
the

1974, resulting

eastern lowland gorillas

is

in

DRC means

are available.'"

Kahuzi-

the loss of

It

has been

of habitat for the

probably the highest tor

any gorilla subspecies, but the lack


the situation

of

in

of clarity

about

that no absolute figures

The fuelwood reserves outside

Kahuzi-Biega have been severely depleted by

wildlife.

DRC, demand

fuelwood by Rwandan

for

refugees affected 105 km'

(1.3 percent) of

Virunga

refugees, so fuelwood collection within the park

an ongoing

is

threat.'"'"'^

National Park by 1997, of which 35 km' had been

completely cleared.'" Since 2001,


Kirolirwe

sector has been

returning to

there by the

cleared

much

the

Disease transmission from

by refugees

Gorillas are susceptible to

DRC from Rwanda, who were

of

settled

Rassemblement Congolais pour

la

Democratie, an armed opposition movement.''''

Another 15 km'

U6

of

land

was

cleared by

Rwandan

as detailed
gorillas to

in

Chapter

humans

as more people

or to

live

in

7;

humans
many human

diseases,

increased exposure of

human

feces

is

occurring

or around the forests, or

enter them more often because they are displaced

Eastern gorilla [Gorilla beringei]

may be

by conflict."" Disease

carried

by park

guards, researchers, tourists, tour guides, loggers,


hunters, or by local people using nearby roads. Data

on the impacts

among

disease

of

eastern gorillas,

particularly outside the Virungas,'" are limited, but

the Ebola virus has not affected eastern gorilla

populations.

Some

eastern gorillas carry parasites includ-

protozoans

ing

nematodes

CapiUana hepatica]," but these


might be

loads

parasite

Cryptosporidium spp.l" and

(e.g.

(e.g.

unrelated

to

human

presence.'^ Mountain gorillas are also susceptible


to

the skin mites that cause scabies or

outbreak

[Sarcoptes spp.l,

an

habituated group

Bwindi

of

mange

which,

in

1996, led to the death

in

an infant male, probably from secondary infection

of scratch
is

in

of

abrasions." " The source of this disease

unknown, but

among
park,

suspected

is

have originated

to

the people and livestock living around the

where

it

is

prevalent." Another outbreak of

in

Bwindi

mites occurred

in

2Q00, but did not result

in

any deaths." Much more seriously, an outbreak

of

pneumonia

in

1988, possibly with an acute viral infection such

in

Rwanda's Volcanoes National Park

as measles as the primary infection, claimed the


but 27 others

gorillas,

of six

lives

successfully.'^"'

'"

The high

rate

of

were treated
infection

181

was new to
against measles was

percent) suggested that the disease

these gorillas." Vaccination

subsequently given

to

65 habituated gorillas from


Alaslair

this

make

While tourism can


to

a vital contribution

conservation by generating funds and through

it
does represent a potential source of
disease" " that could threaten small populations.'^

education,

In

McNeilage

population.'"'"'"

addition, disturbance through contact with hu-

mans may
tibility to

increase stress and thereby suscep-

disease.''

The expansion

exposes more gorillas

to

of gorilla

tourism

diseases that they

may

never have encountered before and against which


they

may have no

natural immunity, while en-

couraging protection of the gorillas from habitat


loss

and hunting. Healthy, fee-paying tourists who

direct targets of hunting but, as discussed above, are


likely to suffer

disproportionately

In

lating

Uganda.

tourism are

place (though not necessarily

in

always obeyed!. These


per day, set a

limit tourist visits to

maximum

and require tourists


distance from

prevention

group size
to

gorillas

of

measures

cm

lands

private

these tourists are


to gorillas.

success

A survey

in

'guarded'

in

tourism with that

were found
immature

likely to

to

of

most circumstances

be on balance beneficial

1981 compared reproductive


gorilla

groups exposed

unguarded groups. The

have

animals.""'

in

smaller proportion

Infants are

not

only

to

latter
of

often

nary intervention

human

is

beings and

in

that

restrictions on

is
In

surround

the

also available at

Uganda, vetericaused by

life-threatening

conditions

to

The Mountain

Rwanda has

human

limited to diseases

that could affect a substantial


a group."

burying

and chasing gorillas

these mountain gorilla centers.

support,

political

mere presence, so

minimum

m."^ Other disease

include

excrement deeper than 30

away from

one hour

of eight tourists,

maintain

deter poachers by

building

Tourists at Mgahinga
Gorilla National Park,

the Virungas and Bwindi, strict rules regu-

parks."" Veterinary assistance

their

are

broken up.

contribute strongly to financing conservation and


to

when groups

number

of gorillas in

Gorilla Veterinary Project

a similar nonintervention policy, with

emergency treatment

to

illnesses

that could threaten the group or population.'^

U7

World Atlas

of Great Apes and their Conservation

Other threats

the reconnected populations are exceeded by the

Gorillas can easily be caught

in

ungulates; this can result

the loss of a hand or

foot.'^'

'"

in

wire snares set for

The three research groups

in

Volcanoes

and research focused on

Conservation

between 1971 and 1998. four

eastern gorillas have been underway for

which had

of

fatal

mam-

mals such as antelopes also wounded many


eastern lowland gorillas

in

Kahuzi-Biega." ' Of

the groups habituated for tourism

in

the

montane

sector of the park, at least one individual per group

had

lost a

hand

in

a snare."

Snares are therefore

the region, mountain gorilla population

illa

size of

populations has given rise

to

mountain gor-

concerns about

inbreeding. However, two studies have suggested


that the Virunga population,

same

size

tion, is likely to

is of

much

the

A comparison

of a

sample

Bwindi gorillas and western lowland gorillas

shows

only minimal reduction of genetic variability

(heterozygosity)

in

the Bwindi gorillas, despite their

nectivity

made

to

however, declining - possibly very

are,

Increased numbers

the Virungas are

in

of

mountain gorillas

probably a direct result of

protection efforts, and are concentrated


two areas. "' These findings indicate
local

commitment and

in

one or

that, with

sufficient investment,

it

is

possible to protect gorilla populations.

The eastern

gorilla is protected by national

legislation in all three of

known populations
not
in

in

range states, and most

protected areas that are

paper parks' (areas protected

or not only,

all,

law, but not

its

live in

practice).

Where park rangers

are present and local residents supportive, gorilla

maintain or restore habitat con-

populations have a good chance of survival. A park

and gene flow between

wherever the risks

some

effort

small population size." Nevertheless, every


should be

in

cases, increasing. Eastern lowland gorilla popu-

be safe from genetic problems for

400 years or more.^' "


of

which

and composition as the Bwindi popula-

in

numbers -

although small - appear to be stable and,

lations

and small

many

met with much

the significant threats associated with warfare

quickly.

isolation

efforts have

success although many problems persist. Despite

the Volcanoes National Park and elsewhere.'"'

The

activities

These prolonged

years.

considered an important threat to eastern gorillas


in

of

gorilla populations,

disease transmission between


Elizabeth

sits with a

A Williamson

rangers work may include monitoring

ment, and community development work. That


parks can

attitudes

is

make

a real difference to local

commitment

illustrated by the

members, who have been known

was

gorilla

populations, patrolling for poachers, law enforce-

gorilla

young female, whose


foot Ijust visiblel

CONSERVATION AND RESEARCH

National Park reported 50 snare injuries to gorillas

consequences. Snares set for medium-sized

A silverback

benefits of expanding the gene pool.

defense

even

of staff

to risk their lives

when pay has

injured by a snare,

in

Virunga National Park,

always been forthcoming. Cooperation and co-

Democratic Republic

of their parks,

ordinated efforts

of

in

park

management

not

involving

the governments of Rwanda, Uganda, and DRC, sup-

the Congo.

ported by researchers and national and international

nongovernmental organizations, have contributed

to

the conservation of the mountain gorilla throughout


its

range, and will continue to do so.

Conservation and research activities

Our growing

understanding

of

biology

gorilla

(including such features as group structure and

dynamics, ranging behavior, habitat requirements,

and population

ways

design

of

political

U8

has contributed

conservation action.

buted indirectly

H'^-'^^-'t;-

densities),

to the selection of protected

in

many

has also contri-

to the raising of global public

awareness, and

Population
helpful

It

in

areas and the

monitoring

of

much-needed

and

funds.

reports are particularly

management, because they provide

Eastern gorilla {Gorilla beringei]

feedback on what
as early warning

working and wliat

is

new

of

gorilla conservation to

as well

not,

is

kinds of threat. This allows

adapt over time,

to

become

including park guards and guides, established a

nongovernmental organization

among

increasingly effective.

The Karisoke Research Center, managed by


tfie

Dian Fossey Gorilla Fund International, has

of gorillas

and

work, as well as studies on social structures,

generated

group dynamics, feeding behavior, habitat use,

national parks, not just

and reproduction.'" Because

making

regular monitoring of mountain

gorillas was 15 months during 1997-1998, a time


when armed conflict prevented personnel from

entering

the

park."'

the

addition,

In

Mountain

gorilla

channeled significant resources

sustained studies of mountain gorillas since 1967.

period without

helped

to

local people.'^'

The revenues created by

These have included long- and short-term census

of Karisoke, the only

that

spread conservation knowledge and reduce conflict

and parts

Chapter

14).

tourism have

into the protection

Box

8.4

money

so

of their habitat Isee

Uganda,

In

the

distributed throughout the system of

is

among

the gorilla parks,

a broad contribution to national

needs and

building political support for gorilla conservation,


albeit at the cost of diluting the

managing

funds available for

populations

gorilla

and

habitats.

Conflicts deter tourists, but during those of the

Gorilla Veterinary Project established a veterinary

1990s, the authorities of gorilla range states (the

center to monitor the health of the gorillas and act

Uganda

in

emergency

situations, including the

removal

of

snares from gorillas and dealing with disease


outbreaks.

Eastern

recently been
gorillas

extended

have more

studies

gorilla

the Bwindi mountain

to

and the eastern lowland

gorillas of Kahuzi-

Biega and elsewhere."'"

establishment

of the

Bwindi Impenetrable

National Park and a buffer zone

1992." The

in

Institute of Tropical Forest Conservation,

Uganda's Mbarara University,


institution of this project.

monitoring program that

the

is

part of

successor

has an active ecological

It

did

what they could

efforts.

The decrease

however,

to

Rwandais du

des Pares Nationaux, and the ICCNl

et

in

to

maintain conservation

revenues from tourism

partially offset in the

Virungas by the contribution

funds and other resources by outside

organizations.

Some

of

the extensive educational

and outreach programs developed prior


conflict also

continued." The continuity

was made
nongovernmental
efforts

Elizabeth

led,

huge enforcement problems. This was

of additional

The Impenetrable Forest Conservation Project


led to the

Wildlife Authority, the Office

Tourisme

to

of

the

these

possible largely by international

organizations

such

as

the

Firewood collection
thie

in

region of the Kahuzi-

Biega National

Parl<,

Democratic Republic of
the Congo.

Willi,3

studying water quality,

is

the impact of forest fires, and forest-gap dynamics.

Other research includes work on barriers

to crop-

and a long-term project on the

raiding by gorillas

ecology, behavior, and population

dynamics

of the

Bwindi mountain gorillas. This research supported

management

the preparation of a

which was updated

plan for the park,

2001 to guide actions for

in

tourism development, biological inventories, and


other measures that are
In

now

in

place.

Kahuzi-Biega, a long-term

community-

based conservation project was established


1985 with the support

the

of

in

German overseas

development agency GTZ, with community-focused

economic development as one of its primary


objectives.'^ " Managers at Kahuzi-Biega and GTZ
developed an emergency plan
things,

collecting

response

to

for,

among

other

and distributing fuelwood

in

the refugee crisis of the late 1990s.

GTZ has also helped

fund gorilla population

to

censuses, including one

in

Kahuzi-Biega that was

also supported by the Wildlife Conservation Society

and others.'"

In

the

same

region,

local

people.

U9

World Atlas

of Great Apes and their Conservation

8A EASTERN GORILLA TOURISM

Box

Research has begun only recently on the


impact on

Rwanda. Uganda, and DRC,

In

tourism

gorilla

and on

of tourist visits,

humans

generates significant revenue, increases public

and

awareness, and has undoubtedly been a motivator

relied

on speculation, extrapolation, and

sense

to evaluate these. Studies of captive gorillas

government commitments

securing

in

protection of gorillas and their habitats.

the

to

should

It

however, be seen as an ideal solution

not,

to the

very specific problem of gorilla conservation, as

exposed

gorillas are

considerable risk through

to

the consequent increased contact with humans.

The
gan

in

cade

first

the 1970s. Far better known,

the program established

is

later in

of the

in

response

Rwanda

in

a de-

to plans to clear a large area

Volcanoes National Park for cattle grazing.

Habitat loss

was viewed as

the greatest threat to the

and

survival of the gorillas, so a carefully planned

well controlled tourism program began as a


of

making the

was

The conservation benefits


include

increased

habituated
patrols.

Daily

surveillance

tourism,

for

With

International

activities

protection

gorilla

of

when

necessary, for

injured gorillas.

began

awareness and concern

has generated funds

and research,

at the

to

recover

for the plight

for conservation

same time enhancing

the profiles of the gorilla range states. The gorilla

was adopted as

a national symbol

and Zaire Inow DRCI, and

is

in

both

Rwandese

the

passport

publicity

and

visas

for

many

made tourism an important earner

visitors
of

of

and

foreign

currency Tourism stimulates the economy, not only


park fees, but also through expenditure on car

accommodation, and restaurant meals.

hire, hotel

People from communities around the parks

gam employment as guides


Uganda, a

Mgahinga
Parks

is

Homsy

warn

to

to

of "the

catastrophic consequences of unconscious gorilla


tourism."'^

Illnesses to which

the

have

gorillas

most dangerous and international

new

carry viruses

to

the

tourists

may

such as novel

region,

strains of influenza. To minimize stress

and risks

to

both gorillas and humans, there are very important


regulations

regarding

maintained

between

maximum number

minimum

of tourists,

distances to be

and

gorillas

people,

the

and the duration

of

their visits, as well as guidelines for appropriate


visitor

behavior

tourist

should never attempt to

still,

to

touch a

Tourism

worse

gorilla.
is

a lucrative business,

which puts

pressures on the gorillas and on park authorities,


leading

some

people to question the continued

justification for gorilla viewing.

The cost

of gorilla-

viewing permits must be set at a level that limits

demand, while maintaining the revenue


to

that

needs

be accrued by the governing authorities. Despite

the

dangers inherent

mechanism

for

in

tourism,

it

provides a

ensuring that national parks and

the gorillas are valued for

many

reasons, and has

certainly contributed to their survival.

Elizabeth A. Williamson

may

or porters, while

in

fixed

proportion of the revenue from

Gorilla

and Bwindi Impenetrable National

contributed to local schools and health

centers through a trust fund.

A tourist and ranger enjoy the


gorilla In

antics of a

young

Vlrunga National Park.


Gordon Miller/IRF

and the advent

organized tourism have attracted

via

have a definite susceptibility

to

diseases, leading

Rwanda

foreigners feature mountain gorillas.

International

common

depicted on bank notes,

stamps, postcards, carvings, and murals. Today


both

human

groups

from poachers, the

population

gorilla

of gorillas

program

this

show them

these studies, conservationists

get closer than the regulation 7 meters or,

monitoring also facilitates rapid

increased

mountain

of

remove snares from

to

and

averted.

and more antipoaching

intervention by veterinarians

example,

means

'pay for themselves,

gorillas

further conversion of park land

gorillas. Prior to

never previously been exposed are potentially the

project to develop gorilla tourism be-

Kahuzi-Biega

though,

150

behavior

gorilla

the risk of disease transmission between

Eastern gorilla [Gorilla berincei)

Programme

Conservation

Gorilla

International

(IGCP) of the African Wildlife Foundation, Fauna

and Flora International, and

WWF-The

Global

run a

number

Conservation Organization. IGCP


of projects.

involved

Is

works with national

and agencies

Institutions

support conservation

and

tias

population censuses, and

In

efforts,

to

strengthen resources,

of national
to 5
is

Other international organizations are also

Inaccessible mountain areas (up

In

home

mountain

to the

gorilla

being produced for Uganda, Rwanda, and DRC.

Comparisons with 1992


the

assessment

computer simulation
aims

images

satellite

changes

of

World Heritage Sites.'

will allow

gorilla habitats in

in

In

a separate exercise, a

of the

Virungas was developed

Dian Fossey Gorilla Fund International.

for the

build capacity.

parks

000 ml that are

movements

to plot the

show

It

through a

of gorillas

habitat preferences,

deduce

collaboration with local organizations. For example,

the carrying capacity of the reserve, monitor

human

the Wildlife Conservation Society has projects

activities (including poaching),

Involved

of the

eastern gorilla conservation, often

In

in

in all

virtual reserve, to

management

eastern gorilla range states:

DRC,

In

It

involved

Is

establishment

In gorilla

park infrastructure, habitat

of

mapping, and exploration


sector
in

in

monitoring, re-

It

of Bwlndl, a

census

of the gorilla population,

Rwanda,

tourism on

In

the

undertaking a study of crop-

is

patterns

around

monitoring,

threats to

success

of

Nations

complementary

is

early warning

ecology,

gorilla

to population
of

and measuring the

and

Scientific

Organization (UNESCO) and

all

World

which alms
Heritage

Images.'^'

to help countries to

Sites via

April 2003. the

In

the

Cultural

the International

space agencies established the 'Open


project,

potential

conservation management. The United

Educational,

use

of

Initiative'

monitor
satellite

European Space Agency

provided significant funding and technical support


for

project with

joint

Environment

the Virungas, and

in

UNESCO

for Gorillas (BEGol.

called

series of

in

in

In

several

Bwindl. These

Rwanda and Uganda, and

by conservation and research groups, with gorilla-

based

programs

tourism

yielding

significant

funding. These gorilla populations are potentially


to

standards

disease and hunting but, by the global

of

great ape conservation,

they are

secure at present.

relatively

Nothing similar can be said about the eastern


lowland

providing

survive

gorillas

parks are managed and otherwise supported by

Volcanoes

the

National Park.

Habitat monitoring

national parks

vulnerable

provides guard support

it

Virungas and

of

and

gorilla behavior;

mountain

small but apparently stable populations

the governments of DRC,

undertaking a biological survey

Is

and studies on the impacts

raiding

lowland

Kahuzi-Biega;

Uganda,

In

the

of

to assist in the

the national parks.'

of

conclusion,

In

and

however, the population status

gorilla,

unknown

which

Is

largely

spread

of

warfare throughout

have been

killed to

following
Its

the

Many may

range.

bushmeat

provide

of

recent

for

armed

displaced people, and miners, and the

factions,

entire population

may have

collapsed as a result. As

the military and political situation remains highly


unstable.

It

Is

very difficult for conservationists to

undertake the fleldwork required

to

circumstances

much

less to

their efforts to

achieve

of

these gorillas,

support local people

In

clarify

humans

sustainable development. The fates of

and

their

needs

for

the

good governance, prosperity,

Build

and peace - are Intertwined with the

maps

wildlife with

which they share

fate of the

their environments.

FURTHER READING
Butynskl, T.M., Kalina,

J.

(1998) Gorilla tourism: a critical look.

In:

Milner-Gulland,

E.J.,

Mace,

R.,

eds. Conservation of

Biological Resources. Blackwell Science, Oxford, pp. 280-300.

Fossey. D. (1972) Vocalisations of the mountain gorilla [Gorilla gorilla beringei]. Animal Behaviour20: 36-53.
Hall, J.S., Saltonstall, K., Inogwabinl. B.I..
gorilla.

Oryx 32

(2):

Omarl.

I.

(1998) Distribution,

abundance and conservation status

Harcourt, A.H. (1986) Gorilla conservation: anatomy of a campaign.

Sustaining Populations. Springer-Verlag,

New

In:

Benirschke,

K., ed..

Primates:

Ttie

Road to

Self-

York. pp. 31-46.

Harcourt, A.H., Stewart. K.J.. Hauser. M. (1993) Functions of wild gorilla


interspecific

of Grauer's

122-130.

close' calls

1.

Repertoire, context, and

comparison. BetiaviourMi. 89-122.

151

World Atlas

of Great Apes and their Conservation

Hart,

Hart,

J.,

T.

(20031 Rules of

Conservation

Homsy,

J.

(19991

in

Practice

i>

engagement
111.

for conservation: lessons

from the Democratic Republic

Congo.

of tfie

http://conbio.org/inpractice/article41RLIL.ctm. Accessed July 13 2004.

Ape Tourism and l-luman Diseases: How Close Should We Get? Report

to the International Gorilla

Conservation Programme. http;//vwvw.mountaingorilla5.org/files/ourworl</Homsy_rev.pdf Accessed February


10 2005.

Hrdy, S.B. (19791 Infanticide

among

animals: a review, classification, and examination of the implications for the

&

reproductive strategies of females. Ethology

Kalpers,
in

J.,

Williamson,

E.A.,

Sociobiology'i: 13-40.

Robbins, M.M., McNeilage,

A.,

Nzamurambaho,

A., Lola, N., Mugiri, G. 120031 Gorillas

the crossfire: population dynamics of the Virunga mountain gorillas over the past three decades. OryxZl

(31:

326-337.
Robbins, M.M., Sicotte,

P.,

Stewart,

eds (2001) Mountain

K.J.,

Gorillas:

Three Decades of Research at Karisolne.

Cambridge University Press, Cambridge, UK.


Schaller, G.B. (19631 The Mountain Gorilla: Ecology

and Behavior

University of Chicago Press, Chicago.

Seyfarth, R.M., Cheney, D.L., Harcourt, A.H., Stewart, K.J. (19941 The acoustic features of gorilla double-grunts and
their relation to behaviour

American Journal of Primatology30\ 31-50.

Stanford, C.B., Nl^urunungi, J.B. (20031 Behavioral ecology of sympatric chimpanzees and gorillas

Impenetrable National Park, Uganda:

diet.

InternationalJournal of PrimatologyU

Taylor, A.B., Goldsmith, M.L., eds (20021 Gorilla Biology:

Multidisciplinary Perspective.

|4|:

Bwindi

in

901-918.

Cambridge University Press,

Cambridge, UK.

van Schaik, C.P, Janson, C.H. (2000) Infanticide by


Watts, D.P. (1995) Post-conflict social events

in

Ivlales.

Cambridge University Press, Cambridge, UK.

wild mountain gorillas iMammalia,

Hommoideal

I.

Social interactions

between opponents. Ethology }00: 139-157.

Weber, A.W. (19931 Primate conservation and ecotourism

in Africa. In:

Potter, C.S.,

Cohen,

J.

I.,

Janczewski,

D.,

eds.

Perspectives on Biodiversity: Case Studies of Genetic Resource Conservation and Development. A/\AS Press,

Washington, DC. pp. 129-150.

Yamagiwa,

Kahekwa,

J.,

J.,

Basabose, A.K. (2003) Intra-specific variation

in

organization

social

of

gorillas:

implications for their social evolution. Primates Ht: 359-369.

Yamagiwa,

J.,

Mwanza,

N.,

Yumoto,

Y.,

Maruhashi,

T,

(1994) Seasonal

change

in

the composition of the diet of eastern

lowland gorillas. Primates 35: 1-14.

MAP SOURCES
Ivlap 8.1
in

Eastern gorilla data are based on the following source, with updates as cited

Chapter

in

the relevant country profiles

16:

Butynski.T.M. (2001) Africa's great apes.


Stevens,

E.F.,

Arluke,

A.,

In:

Beck, B.B., Stoinski,

eds, Great Apes

and Humans: The

T.S.,

Hutchins, M., Maple,

T.L.,

Norton,

B.,

Rowan,

A.,

Ethics of Coexistence. Smithsonian Institution Press,

Washington, DC. pp. 3-56.


For protected area and other data, see 'Using the maps'.

ACKNOWLEDGMENTS
Many thanks

to

Dan Bucknell

(Dian

Fossey Gorilla Fund),

Thomas

Groves (Australian National University], Alexander Harcourt (University


Gorilla

Butynski

International),

Colin

Jose Kalpers (International

Conservation Programme], Michael Wilson IGombe Stream Research Center], and Juichi Yamagiwa (Kyoto

University] for their valuable

comments on

the draft of this chapter

AUTHORS
UNEP World

Sarah Ferriss,

Conservation Monitoring Centre

Martha M. Robbins, Max Planck

Institute for Evolutionary

Anthropology

Elizabeth A. Williamson, University of Stirling

Box

8.1 Juichi

Box

8.2

Box

8.3 Kelly

Yamagiwa, Kyoto University

Martha M. Robbins, Max Planck


J.

Institute for Evolutionary

Stewart, University of California, Davis

Box 8.4 Elizabeth

152

IConservation

of California, Davis],

A.

Williamson, University

of Stirling

Anthropology

Orangutan overview

Chapter 9

Orangutan
overview
Julian Caldecottand Kim

The
is

modern orangutans

lineage that led to

thought

to

have diverged from that

humans about 11
presumably somewhere

African apes and

years ago Imyal,

Asian

McConkey

of the

million
in

the

mainland. Ancestral orangutans then

in-

become the

habited the areas that would

tvtalay

Peninsula of mainland Asia, as well as the islands


of Java,
unit,

Sumatra, and Borneo. This biogeographical

known as Sundaland,'" comprises the lands

above and below the shallow transient seas

theSunda continental

shelf (see

Map

2.11.

out the Cenozoic era, the main land

of

Through-

masses

Sundaland have been joined and separated

in

re-

peatedly from the mainland and from one another


by changing sea levels associated with high-latitude
glaciations and interglacial periods.

The forests
by

general

Sundaland are characterized

of

abundance

Dipterocarpaceae.

These

forests typically have

of

the

Sundaic

family

tree

dipterocarp

poor and irregular

fruit

supply due to their mast fruiting behavior. Within

Sundaland, the abundance

of

dipterocarps and

scarcity of other fruit trees are strong determinants


of

biomass among large-bodied frugivores.

show

taxa

and there
forms

of

tvlany

specific adaptation to these conditions


are,

for

example, specifically Sundaic

macaque [Macaca nemestrina] and

ISus barbatus].^

Much

of

what we know

of

pig

orang-

behavior and ecology suggests a partial

utan

Ian

adaptation along the

same

The Bornean orangutan lineage diverged from


the

Sumatran

mates

1.1-2.3

mya.""" The

for speciation is

the two species

such that

were genetically

it

is

range

of esti-

possible that

isolated

from one

another before being physically separated from

one another^''

If

Singleton/SOCP

lines.

so, speciation

must have occurred

through reproductive isolation: for example, a diver-

gence

of

preferred

mate characteristics

emerging species would lead

in

the two

to a reduction in

gene

transfer between populations.

Populations

[Pongo

abetii]

may

Sumatran

Subadult male Sumatran

orangutan just a few


days after release

in

the

forests adjacent to Bukit

orangutan

Tiga Puluh National

contain remnants of three or

Park, Indonesia.

of

the

153

World Atlas

of Great Apes and their Conservation

shows up
and

populations,''

nized

minor differences

in

in

regional

four

and three subspecies are now recog-

the different parts of Borneo:""'"

Bornean orangutan,

northwest

the

pygmaeus. which
in

microsatellite

in

DNA among

mitochondrial

is

medium

P.

p.

sized and occurs

northern West Kalimantan and Sarawak;

the central Bornean orangutan,

which
in

wurmbii,

P. p.

the largest subspecies and

is

found

is

southern West Kalimantan and Central

Kalimantan; and
the northeast Bornean orangutan, P.

which

is

the smallest and occurs

p.

mono,

Sabah and

in

East Kalimantan.

Both the Bornean and Sumatran orangutans are


large and have obvious sex differences

appear-

in

ance and behavior, with adult males weighing about


75 kg and adult females about iO

Fully dev-

kg.'

eloped adult males have prominent cheek pads


or 'flanges', the development of which

linked to

is

the individual achieving high social status, which

can take as many as 10 years

may

not happen

in all

may

long lived, and

There are a number

of

adulthood and

males.' "' Both species are

reach ^5 years

in

the wild."

of physical differences

between

the two species:

Bornean

the

stouter and

is

stockier,

and

usually has a dark red-brown coat, rather than

the lighter cinnamon fur of the


Singleton/SOCP

Ian

more separate

Secondary lowland
forest in

an old logging

lineages. After the

Sumatran

type, a

new

initial

divergence

orangutans

influx of

from Borneo and the Southeast Asian mainland

area near Bukit Tiga

Puluh National Park

of the

In

thought

to

have entered the area during one

of

is

the

the Bornean has

three types: one linked to the

large,

where orangutans are

Sumatran form; one

linked to the Bornean;

being reintroduced.

with closer affinities to the now-extinct mainland

of

pendulous throat sac and a

figure-eight-shaped

presence
in

the

of a suborbital

distinctive

founded

face

fossa that

is

on

the

lacking

Sumatran species, combined with

forward-facing cheek pads or flanges, while

utan are recognized."

The dispersal

face,

distinctive

developed adult Bornean males have a

fully

may have produced

Sumatran orang-

beard and males a prominent beard and

periods of land bridge connection. Interbreeding

of the

have

moustache;

Sumatra, Indonesia,

orangutan." No subspecies

gracile

around the

fur

little

while Sumatran females

Jambi province,

and one

more

Sumatran;

Bornean orangutans IPongo

pygmaeus] through the Bornean part

of

Sundaland,

Sumatran males have


are covered with

flat

downy

cheek flanges that

hair

where Sundaic ecological conditions are more


pronounced than elsewhere, started

in

the south-

west corner; dispersal was constrained by large


rivers

and high mountain ranges, both

act as barriers to these animals.


later

many

156

became

divided

when

of

which can

ECOLOGY AND DISTRIBUTION


Like all apes, orangutans have simple, globular

stomachs

that cannot ferment food, so they are

The population

limited to eating materials that are not excessively

made

fibrous, toxic, or protected by digestion-inhibitors

climate change

earlier dispersal corridors inaccessible. This

such as tannins.

In

rain-forest

context,

this

Orangutan overview

translates into a diet of sugary, ripe fruit (wliich

range length, and population density

orangutans strongly prefer! and undefended seeds,

to differences

between locations

plus a variety of minor items and 'famine foods'

and continuity

of fruit availability,

sucfi

as

and

insects, flowers,

leaf sfioots,

barl<.

Orangutans are large bodied, so they can tolerate a


certain

amount

material, and will

of mildly toxic

may

often eat soils that

help adsorb and neutralize

in

all

respond

the abundance

seasons, and eco-

system types (both within and between Sumatra


and Borneo). Detailed data on these are

emerging from the few long-term


undertaken,

being

and

are

field

still

studies

supplemented

by

secondary plant metabolites. Being strong and

additional survey information on orangutan popu-

dextrous, orangutans can gain access to edible

lations

items such as seeds and palm hearts that are

both islands, however,

by woody material or

physically well defended

thorns. Their intelligence enables orangutans to

memorize the

locations of cryptic or temporary food

sources, and to use clues - such as the behavior of

opposable toes
adapted
through

allows them to

move

complex three-dimensional environment

orangutans spend most

facility;

in

in

prevent the discovery of

forest ecology.

The preindustrial distribution

was discontinuous

in

of

orangutans

both Borneo and Sumatra.

Orangutan Foundation

orangutan and her

that orangutans are strongly

to arboreality This

with great

time

mean

and

hip joints,

patterns

may

A female Bornean

other animals - to find fruiting trees and lianas.

Long arms, highly mobile

many

and forest composition. Deforestation

offspring. Ttie

mother-infant bond

very close

in

Is

orangutans.

of their

the trees and are deeply familiar with

in

conditions

in

They forage

the canopy

happen

forest unless they

sources of

becomes much more

directed.

the seasonal changes

in

when

their travel

the patchy

in

peaks can

of fruiting

vary with elevation and aspect. These are


attributes of an animal that has

eating niche to the limits

the

large

Orangutans track

production

where the timing

rain forest,

range.'

way through

know where

to

are to be found,

fruit

home

their large

in

a typically zigzag

in

pushed a

all

fruit-

fundamentally rather

in a

fruit-poor environment.

Trees that provide

fruit suitable for

are typically found at higher densities


forest than

Bornean

in

and dynamism

is

much

is

more

in

An

lesser dominance of

the

Sumatra, which are replaced

by other trees that collectively fruit

A number

it

patchiness

the forests of both islands.

in

important feature
dipterocarp trees

Sumatran

and bear

forest,

continuously, although there

orangutans

in

of differences

more

steadily.

between Sumatran and

Bornean orangutan behavior have been attributed


to the different patterns of food supply."

fruit

and

share

to

greater sociability;

orangutans

to

high densities
fruit

those

some

parts of
to

among orangutans
in

eat

behavior from

particular occur at such

in

alone are thought

sociality
to

in

to

it

acquire tool-using

one another Figs

Sumatran

more
among adults, making for
among other things, this allows

orangutans have the opportunity

Sumatra

that these

have enabled greater


in

Sumatra compared

Borneo.' Average home-range

size,

day-

155

World Atlas

of Great Apes and their Conservation

Orangutans were, even then, absent from very large


areas

include the forests


thirds of Sumatra,

River

in

most

southern two

of the

and those between the Rajang

Sarawak and the Padas River

central

in

examples

apparently suitable habitat;

of

in

western Sabah. Prehistoric hunting may have

some

extirpated orangutans from

cave sites

areas

in

now

forests; ancient

lacking orangutans often

may go beyond
"
seasonally abundant foods.' "

exploit

to

residents occupy a

to this interpretation,

defined area over


it

'commuters', or 'wanderers'.'"

of 'residents',

istic

According

Commuters

many

years, but

are seen regularly

in a

particular area

weeks, but vanish and return each year,

for several

repeating this behavior over several years; these

assumed

individuals are

commute between two

to

some

more regular feeding grounds and may follow


waves of fruiting across areas or altitudes. Finally,

locations with abundant orangutans are associated

wanderers are seen very infrequently, sometimes

with cultural influences on people's willingness to

only once, and

contain their bones,

species eaten

along with those of other

by humans.

hunt them; for example,

Aceh province
catchment
long been

An

place

among

hunting taboo has

change

in

is

based on the

close to the edge of an

become

unviable with a

forest composition

example,

of dipterocarps.

tribution

of

breeding

Patchiness

in
in

theories
local

may be

of

orangutan ranging behavior, and interpret


observations

terms

in

all

orangutans having very

of

home

ranges, that

human

extend far beyond the study areas of

observers, and are therefore simply too large to

monitor completely.^^ This would

account for

the dis-

utans that arrive and depart irregularly or at long

populations of orangutans

Both

intervals;

some areas and

ecological absence others.

it

becoming clearer

is

that, the

more

study continues, the

field

likely

longer a
it

is

that

occasional visitors will be seen again.

Whatever the interpretation

correct for different places, with


affecting

an area.

to

'wandering' and 'commuting' behavior by orang-

on which their survival depends.

extirpations

never return

for

favor,

could therefore simply reflect patchiness of the


forests

may

Other authors dispute this classification

large but stable and overlapping

local Iban people.

live

ecological niche that can

the Batang Ai

in

alternative interpretation

theory that orangutans

slight

the strongly Muslim

Sumatra, and

in

Sarawak, where

in
in

in

Furthermore,

or

tions,

and

it

is

of the

observa-

clear that orangutans are not territorial,

that neither sex excludes others

from areas

that they use habitually. Fully adult males, however,

SOCIETY AND PSYCHOLOGY

are intolerant of each other, so they

A mature flanged male

Orangutans

orangutan.

behavior that has been interpreted as character-

are

wide-ranging

animals

with

home ranges

of

may use

they wish to avoid combat. At one study

many as

six

the

other adult males only cautiously,


site,

males ranged independently over

if

as
a

same time, despite ferocious


ensuing when they came into contact with

given area at the


battles

one another"

When young animals

of either

sex

first

gain

independence from their mothers, they often range


widely for a time before settling down,

in

the case of

home range of their


mothers. Subadult males may continue to range

females often close

to

the

widely for a lengthy period and,

if

they do settle, do

so farther away from their mothers.

New

all

'wandering' individuals

seem

to

be adult or older

subadult males; up to 20 percent of these

become established
The
a

individuals

an area are almost always subadult mates, and

in

transition

complex one

in

in a

known

may never

location.

between subadult and adult

male orangutans

is

of both species,

as they exhibit a maturation process known as


bimaturism'.'^ This

156

is

unique

among

the apes and

IS

not yet fully understood. The timing of maturation

is

extremely variable, with puberty beginning any-

Orangutan overview

where between the ages


the

mean

of five

and 16 years, with

somewhat earlier In the Bornean


Sumatran species. There seem to be two

than the

being

pathways

developmental

alternative
'"

thereafter.''

"

Some males

involved

develop certain fea-

tures associated with high testosterone levels, in-

cluding the prominent cheek pads,'

and reach

full

sociosexual maturity sooner than others. These

males are therefore described as

and

flanged',

those of a similar age that have not yet developed

Some males

such features are called 'unflanged'.

remain unflanged and less than


to

mature

fully

for

up

20 years.""

Unflanged males have testosterone levels


Intermediate between those of flanged males and
juveniles.

flanged male

is

thought to have certain

advantages over an unflanged male, notably higher

more secure access

status that gives him

home

established

to

an

range, food sources within

It,

and any receptive females that may be available.


There are costs, however A flanged male

challenges and a higher risk

and high blood levels

of Injury In fighting,

may

of testosterone

themselves reduce lifespan. Mobility


costly for fully developed

body

large

Is

and combative, so there are metabolic

bodied

males due

Is

also

in

more

also

to their larger

These costs may be worth bearing only

size.

if

the individual has a strong chance of gaining high


status and

Hence
flanged

becoming

mate

of choice for

females.

thought that the trigger for becoming

is

It

must be something

relates to the

that

balance between benefits and costs

of high

and

low testosterone levels.


In captivity,

removal

of the

flanged male from

an enclosure will prompt unflanged males

become

flanged.'*

In

the wild,

to

males

flanged

produce regular long calls that certainly inform


females and unflanged males

and status.
of

gap
it

It

seems

flangedness
In

Is

likely,

delayed

of their

whereabouts

therefore, that the onset


In

males

until there Is a

the flanged male population that would

worthwhile for them

to

make

become flanged and


mechanism seems

accept the costs of doing so. The


to

be that the hypothalamus

In

young males, which

regulates testosterone production,

is

affected by

the young males hearing the long calls of flanged

males."

Cyril

at

Ruoso/BOS-USA

around the time

of ovulation.

males

attractive to unflanged

Females are also

A young Bornean

however;

orangutan.

at this time,

unflanged males are occasionally able

them and
In

force copulation

upon them.^""^" Hence,

of offspring,

more

although each successful flanged male

Infants than

does each unflanged male.

Ketambe, northern Sumatra, unflanged males

fathered half the offspring over a 15 year

A male

that

remains unflanged

for a

time

may

catch

both species, unflanged males sire a proportion

sires
In

to

period,''*

with the other half being fathered by a smaller

be subordinate but he avoids combat and metabolic

number

costs and

ern Borneo, subadult males often find receptive

Is

not without mating opportunities, even

though females have a strong preference


adult

males as mates and approach them

for fully

for sex

of flanged

males.

In

Tanjung Puting, south-

females before adult males do, but 86 percent

matlngs

Initiated

by

them are

resisted

of

by the

157

World Atlas

of Great Apes and their Conservation

female.' This
stable

may be two

suggests that there

all

male strategies

the one

at Vi/ork,

linking

Implies a subtle but powerful organizing influence

community

within the

delayed maturation with forced copulation, and the

other linking
intercourse.

maturation with consensual

full

In this

context,

it

is

relevant that early-

CONSERVATION CONCERNS
The most recent estimates

of

total

surviving

developing males do not appear to father late-

numbers

developing sons."

utans are around 7 300 and 57 000 respectively.

The mother-Infant bond


orangutans, but
the

It

gradually

time the apes are

between them

very close

Is

in

weakens with age; by

fully

Interaction

adult.

often limited to glances.'' After

is

Independence, females tend

to stay

near the range

where they were born and maintain

friendly

relationships with local females, which are likely


to

be relatives.

communities

Hence orangutans

that

may

consist

of

live

in

loose

one or more

Sumatran and Bornean orang-

for the

most cases distributed among small and


subpopulatlons

Numbers

of at

of

habitat.""

continue to decline, aggravated by the

secondary trade

now

fragmented islands

In

In

Isolated

in

juvenile orangutans; there are

very few locations where a viable population

250-500 Individuals Inhabits a forest

least

area that

Is

protectable

The

both protected
in

chief

law and potentially

In

practice."

causes

of this decline in population

clusters of related females and the adult male with

are logging, followed by forest

whom

version of forests to farms and plantations [often

they

all

prefer to mate.^^ Researchers have

members
may come

noted that the movements of community

of oil palm, Elaeis guineensis].

are subtly coordinated, and that they

especially

together as a real group on


Individuals
at

In

some

occasions.'"

clusters of closely related females

Suaq Ballmbing, Sumatra, not only share home

potent

in

the

valuable

In

terms

of

timber], and the land

is

more amenable

reproduction; the timing of births

swamp

is

similar within a
If

true, this

below about

timber (especially dipterocarp

and settlement. Even low,

between clusters."

These factors are

lowlands,

500 m, where forests are more accessible and more

range boundaries, but appear to coordinate their

cluster, but different

and the con-

fire,

farming

flat

areas

of

to

farming

deep peat-

forest that are completely unsuitable for

have

been cleared

for

settlement

in

Suherry/SOCP

continuing trade

in

orangutans exacerbates
ttie

decline

populations

in

caused by logging, forest


fire,

and land conversion.

Here, staff of

tlie

Sumatran Orangutan
Conservation

Programme

and an

from the

officer

Indonesian government's

Conservation Department
liave confiscated a

female

orangutan infant from the


village of

Namo Tala

northern Sumatra.

158

in

Orangutan overview

important

Indonesia, reflecting the central-planning failures

former

of ttie

control over

systems

political

tfie

groups

interest

regency levels

management and protection


weak as tfiey were before then,

forest

after 1998,

allowed

recorded

history,

led to the

some

locations; certain local

see advantages

to

in

crucial to extend our understanding of

how

to the increasing

occurring. Our under-

is

in

recent years,

but the extent to which these patterns can

be

extrapolated to regions outside the limited study

and have

sites

protected areas to central

in

is

patterns have been vastly improved

government leaders

Sumatra, which was designated

it

apparently

Most importantly,

standing of the apes' social system and ranging

lost

is

not known.

Almost

studies

Until studies

in

in

Sumatra,

the relatively

Ketambe and Suaq Balimbing.

fruit-rich forests of

200^.

all

example, have been conducted

for

government, such as the Batang Gadis National


Park

sociality,

a relatively fruit-poor habitat.

habitat degradation that

stabilizing

forest conservation

new

started to propose

may be

movements and

in

can better evaluate their response

and the expectation that

from both islands. This situation

determine how Bornean orangutans

orangutans cope with habitat disturbance, so we

and

worst wildfires

lowland forest would soon be

virtually all the

in

provincial

promote very rapid logging

to

and forest clearance. This


in

the

at

to

organize their

regime. Tfie loss of central

have been extended

firm conclusions cannot be

to

other areas,

drawn about whether

WHAT WE DO NOT KNOW

certain

After three decades of field research, large gaps

population, the species, or even the genus; this

remain

in

our understanding

and Sumatran orangutan.


fully

We

of both the

makes

Bornean

the unusual development of male secondary

sexual characteristics. Ranging patterns and social


in

Sumatra, and

it

it

draw firm conclusions about

difficult to

differences between the two species. Lastly, studies

are yet to understand

systems are better understood

are typical of the wider

characteristics

must be extended

to disturbed

understanding

the

of

ecology

areas, to
of

deepen

disturbance,

adaptation, and recovery.

is

FURTHER READING
Bennett, E.L. 11998) The Natural History of Orang-utan. Natural History Publications, Kota Kinabalu, Sabah,

de Boer, L.E.M., ed. 119821 The Orang utan:

Its

Biology and Conservation. Dr W. Junk Publishers. The Hague.

Delgado, R.A., van Schaik, C.P. 120001 The behavioral ecology and conservation
a tale of two islands. Evolutionary Anthropology'}

MacKinnon,
Maple,

T.

J.

|51;

of the

orangutan IPongo pygmaeus]:

201-218.

(1974) The behaviour and ecology of wild orangutans \Pongo pygmaeus]. Animal Behaviour!!: 3-74.

(1980) Orang-utan Behavior, van Nostrand Reinhold

Rijksen, H.D., Meijaard, E. (1999)

Our Vanishing

Company. New

York.

Relative: The Status of Wild Orang-utans at the Close of the

Twentieth Century. Kluwer Academic Publishers, Dordrecht.


Singleton,

I.,

Lacy,

Wich,

R.,

S.,

Byers,

Husson,
0.,

S.,

Stephens,

S.,

Utami Atmoko,

S.,

Leighton, M., Rosen, N., Traylor-Holzer,

eds (2004) Orangutan Population and Habitat

lUCN/SSC Conservation Breeding

Viability

K.,

Assessment: Final Report.

Specialist Group, Apple Valley, Minnesota.

ACKNOWLEDGMENTS
Many thanks

to Ian

Singleton (Sumatran Orangutan Consen/ation Programme], Colin Groves [Australian National

University), Raffaella
for their valuable

Commitante [Cambridge

comments on

University],

and David Woodruff [University

of California,

San Diego)

the draft of this chapter

AUTHORS
Julian Caldecott,

UNEP World

Conservation Monitoring Centre

Kim McConkey, UNEP World Conservation Monitoring Centre

159

World Atlas

of Great Apes and their Conservation

Cyril

160

Ruoso/StiU Pictures

BORNEAN ORANGUTAN [PONGO PYGMAEUS\

Chapter 10

Bornean orangutan
[Pongo pygmaeus]
Kim

pygmaeus

[Pongo

orangutans

Bornean

Linnaeus, 17601 survive

McConkey

306 fragmented

in

orangutans

three subspecies probably

to cross, the

interbreed.*'

and increasingly isolated populations on the


which

island of Borneo,

(Map

10.11."

is

about 7iO 000 km^

The populations

in

area

inhabit forest blocl<s

that are separated by impassable barriers such as

They are concentrated

rivers or areas of cultivation.

Central

in

Kalimantan

and Sabah

llndonesial

West and

[Malaysia], with smaller populations in

Kalimantan and Sarawal< (Malaysia).

East

permanent populations are thought


independent sultanate
though sightings

Brunei

of

No

to exist in the

Darussalam,'"

males have been

of single adult

reported there,' "^ indicating that nomadic indivi-

duals

Next

may wander th