World Atlas of Great Apes and Their Conservation

WORLD ATLAS OF
(iKEATAPtS
AND THEIR CONSERVATION
'
%
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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
Ôj
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
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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
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The contents
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or
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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
.,
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EDITED BY
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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
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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
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
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
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
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
16.17
358
378
Republic of Senegal
15 Lessons learned and the path ahead
354
371

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
distribution
329
Congo
332
Annex: Great Apes Survival Project
GRASP and
Index
433
Partners
439
World Atlas
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
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
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
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
Tlê
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
"
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
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
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
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
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
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
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
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
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
iômo 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
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
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
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
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
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. vîth
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
Caldecott. UNEP World Conservation Monitoring Centre
Martin Jenkins,
28
Box
1.1
Box
1.2 Julian
Martin Jenkins,
(UNEP-WCMC)
for
World Atlas
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
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
Great ape habitats of Sumatra and Borneo
Data sources are provided at the end of
this
chapter
-.-:- '^'-
hi^
I-
'^^^fi
32
<
>
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
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
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
Map
ape habitats
36
2.2 Great
of Africa
this
chapter
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
"
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
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
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
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 lêmoir
(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
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
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
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
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
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
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
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)
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
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
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
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
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
'
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__'^ .âi*
KOUIl
'
Congo
V.a_V
/\\
.1
rA
't
i"
^^' Î
ixJ5*;
vf
T0NG0'Vl|<û^_4r^
DEMOCRATIC REPUBLIC
OF THE CONGO
;
'
r-,,-T
'V,
,_
['
Like
^''
^Â .
^^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^Âr
*^
ANGOLA
10"S
10E
20'E
55
World Atlas
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
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
of
the Oban
Country
Bating Iproposed Biosphere Reservel'""
Mall
western
Bossou, near the Nimba fôuntains"^^'
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""""
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'"
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
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
îy
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
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
(^ôvement 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
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
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
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
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
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
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
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
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
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
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
[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."

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
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
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
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
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
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
captive conditions
1999
animals stressed
release cage
boat;
5 I2M, 3FI
and panicked by
captive conditions
2000
4|1M,3FI
floating
boat;
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
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
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
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 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
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
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
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
in
law enforcement;
and
for
by corruption
chimpanzees remains
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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
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Struhsaker,
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Matama,
Goodall,
H.,
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in
213-228.
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45-54.
Reynolds.
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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
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
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
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
Bonobo
distribution
Da'.i ~-n,,rrrs
are provided at thp pnd of
this
chapter
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

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
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
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
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-
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
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,
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
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
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
Vîldlife
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
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
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
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
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
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
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:
T.S.,
T.L.,
Norton,
B.,
Rowan,
A.,

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
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
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

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
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
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
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
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
T.S.,
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.,
tôuntain 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.

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
Julian Caldecott,
Sarah Ferriss,
103
World Atlas
TJ. Rich/natureplxom
106
Western gorilla [Gorilla
gorilla)
Chapter 7
Western
gorilla
[Gorilla gorilla]
Sarah Ferriss
western
The
18^7)
gorilla
Savage,
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
Western lowland
""
Box
gorilla distribution (see
'ôE
~ "Ty
7.1 for
Cross River
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
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
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
'
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
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.
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
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
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
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
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
(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

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
A-ta^îzvi
.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.
''
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
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
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
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
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
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
in
the population
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
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
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
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"
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
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
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.
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
{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
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.
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
127
World Atlas
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
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
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
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

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-
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
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
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
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
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
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
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
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
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
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
(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
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
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
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
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
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
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
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
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
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
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
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

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
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
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.
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<,
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
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
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.

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., Nlûrunungi, 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:
|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.
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,
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:
eds, Great Apes
and Humans: The
T.S.,
T.L.,
Norton,
B.,
Rowan,
A.,
Ethics of Coexistence. Smithsonian Institution Press,

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
AUTHORS
UNEP World
Sarah Ferriss,
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
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
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,
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
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-
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)
AUTHORS
Julian Caldecott,
UNEP World
Kim McConkey, UNEP World Conservation Monitoring Centre
159
World Atlas
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 through

to
Brunei
is
from the Rajang River

in
Sabah. This
the forests on occasion.
a large area of recorded absence,
may
in
Sarawak
Padas River
to the
reflect local extinctions
by hunting pressure
in
caused
the past,'^^ an unsuitable
Bornean orangutan populations are divided
medium
Pongo pygmaeus
least three subspecies.
pygmaeus
(the
northwest Bornean orangutan!
sized and occurs
Sarawak. The largest subspecies,
P. p.
central Bornean orangutan],
is
found
Kalimantan; the smallest, P.
morio
p.
Bornean orangutan) occurs

Kalimantan. There
is
is
Indonesian
(the northeast
may
also be distinct
in
South Kalimantan,
province that forms the
eastern part of Borneo.
south-
Rivers form the main
barriers between the subspecies.
Borneo, where
Central
yet to be fully accepted.^'
Orangutans no longer occur

the
in
evidence that the Sabah and
East Kalimantan orangutans
subspecies, but this
wurmbiilthe
Sabah and East
in
In
Bornean orangutans have
the middle of
the rivers are sufficiently
small
for
strongly expressed
preference for large, succulent
energy content
when
of
in
they occur
such
fruit
with a high-
fruit
the form of sugars, particularly

large crops.'*
in
influences virtually
Bornean orangutan's
The
availability
aspects
all
of a
ranging patterns, social
life: its
"'"
behavior, timing of reproduction, and health."
makes up over
Fruit
10.11,
50 percent of the diet (Table
but orangutans are opportunistic foragers
rather than absolute frugivores. Dietary composition
may
therefore change markedly with time of
year," as orangutans
times
make use
because
of 'famine foods' in
opportunism and dietary
of shortage. This
flexibility is
essential to a large-bodied forest animal
its
amounts
of
mature leaves
simple, globular stomach lacks fer-
mentation chambers.
Bornean forests are renowned
is
West Kalimantan and
in
Feeding, foraging, and ecological strategy
that cannot digest large
environment, or a combination of both.
into at
for
irregular food supply for fruit-eating and
their
seed-
eating animals. They support a low frugivore bio-
mass compared
to that
supported by other tropical

'"'
moist forests, such as those

is
largely a result of the
fruiting tree family, the
dominated
by
in Africa.'"'
abundance
"' "' '"
This
one mast
of
Dipterocarpaceae." Forests
dipterocarps
are
seen
also
Sumatra and Malaya, but are more widespread

Borneo. These trees produce huge crops
chemically
undefended,
oil-rich
of
in
in
winged,
seeds during
years apart, with
many
different dipterocarp species participating in
each
masts' that occur one
mast. Masting
is
to six
complex form
of tree
behavior
161
World Atlas
Map
162
10.1
Bornean orangutan distribution
this
chapter
BORNEAN ORANGUTAN [PONCO PYGMAEUS]
that
linked to drought, which
is
by the
influenced
cycle/^' '"
and
As trees
El
of
response
fruit in
Borneo
in
is
strongly
Nino Southern Oscillation
many
other families also flower
masting events often
to drought,
involve a large proportion of all trees in the forest.
Frugivores
in
this irregular food
Borneo are adapted

supply They
masts and starved
during
at
cope with
to
Several authors have concluded that orangutans
sources."
direct routes
remember
advantage
trees enable
mosaic'
of a 'phonological
long legs and
its
in
it
to
which
take
fruit-
peaks move around over large areas."
to
scheduled food
have
between them;
to
They gorge them-
little diffi-
and can take
suggests that they
this
food plant seasonality and locations.'*"'
Observations also suggest that orangutans
can interpret clues regarding the location
of fruit
sources, such as the strong smell of durian fruits
iDunospp., Bombacaceael, the noisy interactions
of
mass movement
of
other feeding animals, and the
hornbills IBuceros spp.) and flying foxes
Orangutan strategy has some similarities

that of bearded pigs IBox 10.1).
differently
Orangutans appear
culty locating available fruit sources,
other times. This
very mobile;
swim and even climb
ability to
ing
is
'
con-
of elevations, topographical features,
and forest types, each with
risk being satiated
For example, the Bornean bearded pig ISus
barbatus barbatus]
range
tain a
situation has favored mobile species with flexible

diets.
may
track food supply across a landscape that
(fruit bats,
Pteropus spp.l toward new feeding grounds.'^'
The congregation
of
animals around
'"
a tree with ripe
some
selves during mast fruiting, increasing their calorie
figs, for
intake by 50-70 percent, eating only fruit (including
distance and often attracts orangutans.'" These
When
behaviors suggest a significant capacity for making
dipterocarp seedsl while this
explosions
exploited
main
in
in
is
abundant."'^"
insect populations occur, these are
in
a similar way.'"
Orangutans may
re-
the vicinity of a fruiting tree for several days
until the
crop
is
exhausted. The orangutans store
during these times, which helps
them
fat
to survive for
based on a detailed spatial

memory,"' '" and are suggestive of how a largeintelligent decisions
bodied, semisolitary, arboreal
ceed
is in
in
is
to
follow a
steady zigzag pattern through the forest, feeding on

different sources."^ Individual
orangutans may
be familiar with an area of over 30 km'
Table 10.1 Diet and range
in
in
extent.'"
ling figs, are
mast
fruiting
pattern of
can sucfruit
many
to the
to
the pattern of
annual-to-biannual fruiting
other tree species." Fig trees
may do
fruit
so two or three times a
three Bornean orangutan populations
Kalimantan""
of study years
an exception both
and
abundantly, and
Tanjung Puting
Number
mammal
ecosystem where
a living in an
Figs [Ficus spp., Moraceae), especially strang-
Their usual foraging strategy
many
making
short and unpredictable supply
weeks afterwards.
a few
example, can be seen or heard for
Kutai
Kalimantan"
Kutai
Kalimantan''
Kutai
Kalimantan"
Ulu
Segama
Sabah''
Diet
No. of types of food eaten
317
species eaten
169
No. of
fruit
Proportion of diet (percent)'

61 (16-92)
54(13-89)
61
53(10-97)
Young leaves
15(0-40)
29 15-57)
25
36 (7-73)
Flower
4(0-41)
2(0-11)
Bark
11 (0-47)
14 10-67)
13
16(0-37)
Insects
4 (0-27)
2 (0-8)
Fruit
10-3)
Ranging
Female home range (km')
5.0-6.0
0.4-0.6
0.5
0.6-3.0
Male home range (km')
6.0
1,0
4.0-8.0
Female day range (m)
710
305
500
Male day range (m)
850
305
500
Mean
figures, followed in parentfieses by tfie montfity
range observed
163
World Atlas
such circumstances, they consume
In
fruit.''
large quantities of bark (37 percent of diet
one
in
study!," leaves, gingers (Zingiberaceael, and stems.'"
Orangutans also consume the growth layer under

the bark (mainly of Ficus trees); their teeth
adapted
being
signs of
orangutans studied
in
to
this
task.'^
show
Several
Kutai, East Kalimantan,
were
able to persist for half a decade by consuming
bark and vegetable matter, after their forest was

devastated by
of
1982-1983.'
fire in
some areas
In
'
Borneo, the young leaves, top shoot, and inner
meristematic tissue
of the
mature
stem
leaf
of the
palm Borassodendron borneensis form extremely

important food sources during periods of
fruit
scarcity Local people insist that orangutans can
survive only
areas where this species
in
is
fairly
abundant.""'"-'"
No confirmed observations
been made
An orangutan
eating tree
for the
was observed
infant
of carnivory
biting off a rat's
head before
bark, a particularly
playing with
important food source
year old was seen
when
a bird before discarding the body." Bird
fruits are scarce.
also
each tree on a different
year, with
peal<s
in
home range
figs at all times.'''
other
fruit
present
in
to
for
be likely to contain
Orangutans
preference'^ but,
typically eat
where
figs
are
and there are few alternatives, they
comprise an important part
orangutan
of the
Orangutan populations also occur
swamp
as the
There are
abundance, but these are weak enough
an orangutan's
some
cycle.
areas such
forests of Tanjung Puling National
Park that are virtually devoid

staples
forests,
in
diet.
include
iTetrameristaceael, which
of fig trees." In
Tetramerista
fruits
these
glabra
throughout the
body," and a formerly captive two

to bite off
and eat the head
of
eggs may
be eaten."' Leaves, young shoots, flowers
(notably Madtiuca spp., Sapotaceael,
wood
pith,
mushrooms,
honey, insects (leafhoppers, crickets,
bees, ants, and termites], and mineral-rich soils are

'" '
certainly eaten."' '"
These soils sometimes
have high concentrations

to
of kaolin,
which may help
adsorb and neutralize the large amounts
tannins and
consumed
selective
of
other secondary plant metabolites

the
Bornean vegetation, despite
feeding.
Orangutans have been seen
in
eating sections of the tubes of soil deposited along

tree trunks by termites,
to the
ground
to eat
and may also descend
clumps
of earth.
They
visit
Xanthophyltum rufum IXanthophyllaceae);
'mineral
licks'
Gironniera nervosa lUlmaceae); Lithocarpus spp.
trations
of
(Fagaceael; and Nephelium spp. [Sapindaceael.
potassium, and calcium!; these are visited by a wide
year;
In
the drier alluvial valleys, Ficus spp., Agtaia spp.
Baccaurea
(Meliaceael;
spp.;
such as caves with high concen-
important
minerals
variety of animals, including deer
and Mallotus spp.
(e.g.
sodium,
and gibbons."'
Male and female nutritional needs are some-
(both Euphorbiaceael are all important dietary

sources for orangutans." " Several tree genera fa-
what
vored by orangutans
(e.g.
time on the ground where termites occur." Female
produce
of
also
importance
to
fruits
Durio, Agtaia, Nephelium]
dietary and
commercial
humans.
Other orangutan adaptations
to fluctuations
in
food availability are their
to
unusual food sources. Outside masting periods,
flexibility
and resilience
orangutans regularly consume unripe
and other
fruit,
seeds,
items.'^ During periods of food stress,
less than 16 percent of the orangutan diet
16/>
its
have
Bornean orangutan, although an
may be
different.
Males eat more young leaves, bark,
and termites than females do; they also spend more
orangutans eat a more varied
diet in
consume fewer
males."" Females
calories, than
therefore store less
more
fat in
times
generaL but
of plenty,
and are
affected by periods of shortage."
Ranging behavior
Bornean orangutans are wide-ranging animals
that
occupy potentially very large home ranges; being
BORNEAN ORANGUTAN iPONGO PYCMAEUS]
Box
ADAPTATIONS OF BEARDED PIGS TO

DIPTEROCARP FORESTS"'""
10.1
LIFE IN
gestation length 190-120 days], permitting up to two
Bearded pigs [Sus barbatus] consume
roots, fungi,
small animals, turtle eggs, carrion, and items from

at least
50 genera
and breeding, and
growtfi, fattening,
seeds
and
of
supply controls
of plants. Fruit
tfie
oil-ricfi
dipterocarps \Shorea, Dipterocarpus spp.,
otfiersl,
oaks [Lithocarpus. Quercus spp.
I,
tant.
Highland oak forests produce
fruit regularly,
and pregnancy (10-20 months);
rut
conversion of dietary
potentially high
body
fat to
growth rates; very
efficient
variable but
fat;
group
flexible
sizes 110s to 100s); synchronization of birth peak
with fruit
and travel-adapted features such as
fall;
swimming
long legs and
ability
The synchronized
and
chestnuts [Castanopsis spp. are especially impor-
per year; variable but potentially early age at
litters
first
birth
prompted by the
a rut,
peak
falling of
from the forest canopy
achieved by
is
massed flowers
Male
(the 'confetti effect').
and are important as predictable food sources,
bearded pigs bombard the females with olfactory
while dipterocarp forests provide large supplies of
and/or pheromonal signals from their urine.""
food
only
fruiting
is
Irregularly
well
of
Dipterocarp
masting.
documented
many
the seeds of
by
in
Sarawak because
these trees are exported
commercially, and records have been kept for
many
decades. Since 1899, virtually no dipterocarp seeds
were exported
in
in
one
small amounts
third of years,
most other years, and more than
some
Episodes
years.
occur every three
of
heavy
to five years.
1945-1988 were four pairs
000 tons
in
usually
fruiting
Sadly for the male,

rut,
he must go
pating
and
1986-1987).
utans delaying
events were correlated with

linked to El Nino
The bearded
of their entire lifetime
may
during
droughts
repeated
Southern Oscillation phenomena.
pig
population rose dramatically
1954, 1959, 1983,
and 1987, but
1945-1988,
in
suggesting
population growth occurred
worth the
is
cess
these
in
in
no other year
that
explosive
response
to
sus-
risk of death.
In
hundreds
sire
on one
rut,
during which
of piglets.
Malay Peninsula, there
the
take advantage of predictable fruiting
wood [Dryobalanops aromatica]
in
forests,
the eruptions
in
bearded
Pfeffer described
pig
numbers,
in
Kalimantan over distances
of
250-650 km;'"
large average
to 12,
litter size
Iwhich can range from three
depending on the mother
pig's size); short
Caldecott tracked
In
of
Sarawak
8-22 km/month, migrations that were
sustained over four or more months." Large-scale
movements
of
populations of bearded pig have also
been reported from the Malay Peninsula.^^'-^^'^'
A bearded
in
the
herds of bearded pigs
19B0s,
at a rate of
the 1950s,
annual population movements
moving through the upper Baram area

allow this response Include
camphor-
which have
since been felled for their valuable timber. During
consecutive years.
may
historical
is
evidence that bearded pigs migrated regularly to
tained availability of dipterocarp seeds over two
Features that
9] until
Male bearded pigs may stake the reproductive suc-
they
1980s,
delay partici-
factors suggests that the
of
success
likelihood of
may
maturation; see Chapter
full
some combination
years with heavy
During the
achieve a successful
[analogous to unflanged male orang-
in a rut
Exceptions during
of
to
acute urinary retention, which
can destroy his kidneys. Males
dipterocarp fruiting 11953-1954, 1958-1959, 19821983,
order
in
Into
"
During these, the pigs were described as moving
pig.
Cede Prudente
consistently
in
one
direction, in scattered or
densed herds, over either

this
was observed
a broad or a
narrow
confront;
to persist over a period of days,
weeks, or months. The animals were variously

described as being
good, poor, or very poor
in
physical condition. They
were sometimes accom-
panied by piglets and sometimes not, and regularly
swam
bays,
lation
across
rivers,
and even out
was thought
sometimes across coastal
to sea. in
some
cases, the popu-
to retrace its route later,
follow a circular course to
its
or to
point of origin.
Julian Caldecott
165
World Atlas
very loose, dispersed communities with
utans
live in
little
interaction
compared
most great apes.
to
Mating opportunities are restricted by population

density, as well as by female receptivity; this, in
turn,
thought
IS
be related
to
food supply."
to
Population densities vary across Borneo, with

relatively
dense and social
much more
frequently than do
from the
individuals
Kutai area mating
those from Gunung Palung National Park (West
same
Kalimantan!. The
pattern
seen
is
for all
other aspects of sociality, including aggressive
which are regularly observed amongst
interactions,
Gunung Palung.The
the Kutai apes but seldom at
larger and
Bornean orangutans have not been seen
fact that
Flanged males are
more
to
use tools
in
aggressive than
Sumatran
unflanged males, as
consequence
well as being
more
the wild, unlike their
has been interpreted
relatives,
of the
social
be a
to
lack of opportunity for indi-
viduals to learn from one another'"
more
Bornean orangutans spend most
attractive to receptive
alone or
females.
of their
time
mother-offspring binary units. They
in
associate loosely with others at times and are no

also slow moving, their daily travel distances are
small relative
the size of their
to
ranges vary considerably

sites Isee Table 10.11.
in
home
range.
Home
area between study
They are not defended and
overlap extensively, although fully adult males are

intolerant of each other
The ranging pattern adop-
doubt familiar with the individuals that use their
home ranges
to
having been observed
fairly regularly,
coordinate their travel."' Familiarity and related-
ness presumably go together,
females,
at least for
as they tend to settle near their mothers." Groups

of
two or more individuals, excluding mother-infant
formed 19 percent
observations atTanjung
ted by a particular individual appears to be deter-
pairs,
mined by
Puling" and 20 percent of observations
position,
a combination of food availability, social
and reproductive
Females use
condition.'''
a smaller area of forest than do males, usually
the range of 0.5-5.0 l<m', especially

restricted
in
sistently at least
those
movements
their
The home ranges
infants.'
two
to three
by
if
in
they are
times larger than
High-status or dominant adult
of females.
males, however, appear to maintain a relatively
home range
small
during their period of domi-
nance, within which they attempt to monopolize
access
females."" Individuals
to receptive
new
to
area are almost always subadult males, and
seems
certain that
Segama.'^ Associations or
at large fruit sources, but
travel
males are the dispersing
an
it
sex,
some
fully adult
social orangutans,
their
gregarious.
may
interest in
of both
percent of
play together, even
each
other.*'
genders can also be
Tanjung Puling, adolescent
In
females spend about ^5 percent

groups, compared to
form parties
likely to
spending 91
mothers show no
Subadult orangutans
quite
may also
Bornean males are the
their time alone." Juveniles
when
individuals
Ulu
occur
between food sources together"
than males;" '"

least
at
'parties' typically
Females are much more
dependent
males are con-
of
of
of their
time
in
percent for adult females."
Subadult males spend about 41 percent of their

groups, of which 83 percent
with solitary males being encountered at times far
time
from locations that support breeding populations
females, and only 3 percent spent exclusively with
(for
example
at
high
altitudes
in
Sabah and
in
males."
In
spent with
the presence of adult females, adult
males do chase
Sarawak)."'"'
is
off
latter are able to
subadult males, but usually the
move
quietly
away"
Party formation
The scarcity
most
166
of
of large, reliable food
Borneo
is
thought
to
sources across
be the key factor that
Relations between males
The large body
size of fully adult males,
reduces the opportunity for social interaction
consequent need
among
ripe fruit,
orangutans.'" Typically,
Bornean orang-
to
consume
and
their
large quantities of
makes resource competition
inevitable
BORNEAN ORANGUTAN [PONGO PYGMAEUS]
same
overlap with their own; others
may
defend territories and aggregations
of
and mate opportunistically
the
areas with an abundance
of
other males, sometimes by forcing copulation.
between males
actively
the
in
males can occur
in
area.
Males do not
may be
resources, but adult males are aggressive towards
Unflanged males
each other
male and spend most
close range;
at
sometimes with
battles
'"" Combat
fatal results."
as adult males tend
to avoid
can ensue,
rare,
is
each other
in
more mobile
be
home ranges
of
tolerated by a flanged
time within his range,
of their
may wander in search of food and females.

Such males may consort with females but most
or they
copulations that follow are forced. The implication
of
Female orangutans reach maturity between the
with
ages
of
1 1
and 15 years" and reproduce about every
eight years,"
''
depending on ecological conditions.
The processes
of
male maturation
one that involves some males becoming
fully
mature, or 'flanged', earlier than others that remain
sometimes
unflanged',
Chapter
91.
for
many years
Flanged males have larger bodies, are
more aggressive and dominant over the

are the preferred
actively
Isee
mates
others, and
of adult females.
The
latter
seek and frequently obtain attention from
males as they
represent
little
Orangutan pregnancy
long,
dependent
position until
occur during periods
when female estrogen
of
mast
In
is
fruit
at
half
a side-ventral
in
several years old. By 11 months,
Mothers may play with
them
and a
young survive their
own
food,'' but
may
years of age."^
their young,
and sometimes
directly.
seven
its
five or six
Offspring
to 10
leave the maternal
years
home
become
range."'
fully
when
they
among
the
of age,
"^
may
and
is
fruiting,
Vocal behavior
The long
call of the
male orangutan
is
strangest and most penetrating sounds heard
levels are highest."
Mating can take two basic forms.
it
continue to suckle until
may
likely to
of
While traveling, a mother
infancy.
the young orangutan can find
neal swellings of the chimpanzee (although vulval
most
lasts for eight
orangutan carries her offspring
independent
cycle of mating,"^'
competitive threat.
months.""^ About 90 percent
not display an obvious physical signal like the peri-
more than one
mate
tolerate the presence of unflanged
feed
require
that females prefer to
males; these
flanged males around the time of ovulation," but do
swellings occur during pregnancy!. Conception
is
high-status, fully developed
complex
is
these observations
the
first,
nature.
These
in
calls are given by the highest-ranking
the female initiates contact with a flanged male,
males three or four times a day and are often
which then actively
accompanied by branch-shaking displays and
by
means
Intercourse
solicits
and a penile
posturing
of
sexual intercourse
display.'"'^
consensual and often occurs with
is
the female 'on
top';^'-
same male during
"^
it
may be
repeated with the
consortship lasting several
consecutive days, or the female
may mate
more than one male around the time

The second form
of
bristling of hair"
arrives at a
new
the direction
Cyril
in
Long
calls are given
location; as
when
orangutans often
which they are
male
call in
traveling, their calls
Orangutan infants have

a long period of
dependency
relative to
that of other great apes.
Ruoso/B05-U5A
with
of ovulation.
orangutan mating involves
forced copulation by unflanged or, less often, by
flanged males that are strangers to the female."

either case, juveniles are often intolerant of
mating with their mothers and

male."^
Mating
Tanjung Puting
may
behavior by unflanged
attack the
males
in
National Park has been docu-
mented by Birute Galdikas." Here, 86 percent

matings
In
males
initiated by
of
unflanged males were resisted.
Unflanged males often locate receptive females

first,
but are soon displaced by flanged males.
An
unflanged male does not attempt copulation with

a pregnant female or
when
a flanged
male
is
in
the vicinity.
Some flanged males
maintain a circumscribed
range and mate with the females whose ranges
167
World Atlas
and bananas was spat

slowly."'
that
then eaten
into the vessel,
These observations are taken as evidence
Bornean orangutans are capable
and transmitting
of using tools
behavior culturally, as do
this
Sumatran orangutans, but
that they
meet each
other too infrequently for this capacity to be highly
developed
in
the wild.
Nest building
Every night, just before sunset, orangutans con-
These are springy platforms,
struct night nests.
formed by bending and weaving small branches.
They are often covered with loose twigs and
may
branches. An existing nest
be used,
if
situated
close to a prime feeding tree. Orangutans usually
remain
night.
midmorning, although
their nests until
in
some
occasionally
move during
individuals
Bornean orangutans often nest close
last food tree visited that day,
to
the
the
and feed there again
the next morning; they frequently return to the night

nest for a late-morning rest. Orangutan nests are
considerably easier
see
to
in
the forest canopy than
are the apes themselves. They can be distinguished

This orangutan nest
few days
old:
leaves are
color.
most
still
is
of the
green
in
may
allow males
to
reduce the chance
space themselves out and
of violent interactions.
Long
calls
are often given after associations with females and

are
more frequent
in
areas where there are few
consortships underway." They
may have
attracting females, or at least
alerting
male's presence,
to
have
in
from those
easily (at least
fore be
used
of
other
Natural enemies
their heads,^ but have not
to tal<e shelter
sense
been observed
to
use
using one object to
of
another Captive Bornean orangutans,
however, learn tool-using behavior quickly from

others, including
humans;
in
zoos, both
Sumatran
and Bornean orangutans can surpass chimpanzees

speed
of invention
and learning, and
Rescued orangutans
creativity'"
at
Camp
in
human
humans
riding
in
canoes,
hammering
nails,
in
many
'brushing'
activities,
teeth,
probably
at
in
is
no evidence
of
Borneo, but juveniles
threat from
clouded leopards
[Neofelis nebulosa], pythons [Python reticulatus],
and black eagles [Ictinaetus malayensis].'' Iban

folklore
has accounts
diles but the
of
orangutans fighting croco-
two species are unlikely
when an orangutan
to
falls into a river
Some juvenile orangutans

have been
killed by wild
Response
to disturbance
meet except
by accident.'^
undergoing rehabilitation
bearded pigs."
logs,
shooting blowguns, and washing
Orangutan
densities
are
reduced
by
habitat
disturbance,"' '" with the extent of this depending
on the severity
showed
of
damage
to the forest.
One study
that densities differed by only 7 percent
to
between disturbed and undisturbed lowland dipter-
use leaves as a feeding vessel. They carefully
ocarp forests, but by 21 percent between disturbed
laundry.'"
In
Sabah, rehabilitated apes learned
selected leaves and arranged

able to hold semifluid material.
168
including
sawing
are
there
activity,
their
Leakey,
Tanjung Puting National Park, have been observed

imitate
orangutans;
quickly for nests.'
development
full
9|.
strict
of
seem
the
Bornean orangutans have been seen
to
assess the density
addition to the effect they
from the sun or rain by holding twigs or leaves over
their
to
helicopters can be used to survey large areas very
predation on orangutans
tools in the
of giant
to the
in
Tool use
in
from the ground) from those
squirrels [Ratufa spp.l. Nest observations can there-
Aside from
influence
other animals, such as
of
them
in inhibiting
males (see Chapter
a role in
readily
sunbears {Helarctos malayanus], although less
them
into a vessel
A mouthful
of
milk
and undisturbed peat-swamp forests
in
Gunung
Palung National Park. The authors suggested that
BORNEAN ORANGUTAN [PONGO PYGMAEUS]
Box 10.2 ORANGUTANS
DEGRADED
IN
tangan floodplain have been sub)ect to increasing
HABITATS
pressure from commercial logging and agriculture.
As a
Sabah
more than
IMalaysial harbors
10 000 orang-
climax (mature!
result,
have been
forests
from the towering, highly struc-
greatly transformed
utans with most
of
outside primary
tured ecosystems of the past. These forests
now
forest reserves,
forests that have been selectively
have a low stature, lack a clearly recognizable
divi-
in
these
living
logged,* Rather than regarding orangutans living
in
secondary forests as ultimately doomed, the French

organization Hutan, together with the
Department, conducted the
first
Sabah
Wildlife
detailed long-term
study on
how orangutans cope with changes
natural
habitat,
enhance
their prospects of long-term survival.
seeking
in their
solutions
realistic
to
The
sion of canopy layers,
density of 332 trees
over 10
up
1998
in
the Kinabatangan floodplain
in
in
trees that have a diameter
at breast height]
per hectare. There are
very few large trees, small basal area, large gaps
these attributes are characteristic of disturbed

habitat,^
The emergent trees belong

Dipterocarpaceae,
to
Although
fragmented forests that
trees are never taller than 35 m,
wildlife diversity
harbors remarkable
and abundance, including one
the largest orangutan populations
Since the mid-195Gs,
all
lowland dipterocarp type!
of
in
of
Malaysia,^"
forests (mainly mixed
the
Lower Kinaba-
this
is
comparable
the heights of 70
to
(vioraceae.
primary
in
more reached
or
forests.'" Fast-growing
forest, the
contrast to
in
primary
and light-demanding
neer species that are tolerant
show
the families
Leguminosae. and
eastern Sabah, a corridor of highly degraded and

still
in
the canopy, and a high degree of soil disruption;
Kinabatangan Orangutan Conservation Project was

set
cm
and have an overall low stem
(re,
of dry conditions
pio-
and
opportunistic life-history characteristics are

continued overleaf
Plant families most often
consumed by orangutans
Leaves
Fruits
Feeding time allocated

Fruit family
to
item type as percent of total^
66
as percent of food type
Moraceae
27
13
Fagaceae (beechi
12
<1
<1
Anacardiaceae (mangel
(figl
10
<1
<1
Ebenaceae (ebonyl
18
<1
Rubiaceae
<1
11
Polygalaceae (milkwort!
<1
<1
Sterculiaceae (cocoa!
<1
32
Woody climbers
13
ib
28
32
15
25
(coffeel
(except Moraceae)
Other families
Bark
22
During
700 fiours
this reflected
peat-swamp
illegal
much
of
in
Sukau study
higher disturbance rates
forest.'^
logging
feeding observations at the
in
the
Between 1999 and 2004, the
and around Gunung Palung
National Park accelerated, with removal of large
numbers
of
fruit
trees,
halting
research
in
into
areas
of
crowding led
shifts in
to stress,
and a decrease
In
the
Only limited assessments are available of the
cope with such degradation."
forests of the western
Central Kalimantan,
illegal
!n
the peat-
Sebangau catchment,
logging
operations
orangutan distribution, often
suboptimal habitat. The resultant over-
in
increased juvenile mortality,
fecundity."
Kinabatangan Wildlife Sanctuary
Sabah. a research
behavioral response of orangutans to and their
swamp
caused large
the
park.'"'
ability to
site.
site
in
has been established by
the Kinabatangan Orangutan Conservation Project

to
investigate
orangutan populations
forest, initial results indicate the
high population density." although this
temporary consequence
of
the
in
presence
logged
of a very
may
be a
concentration
of
169
World Atlas
common
these forests:
in
Macaranga
Euptiorbiaceae
Rubiaceae,
spp.l,
Moraceae. creepers [Spatholobus
The
spp.l.
Meremia
spp.,
and grasses [Paspalum
spp. and Uncaria spp.l.
Imperata
le.g.
Stercutiaceae,
diversity
of
spp-.
these disturbed
Kinabatangan forests remains high by gtobal standards, with
minimum
of
056 tree species
belonging to 129 families and 512 genera.'

Today, the composition and structure of these
forests
is
a mosaic of different types of habitat,
at different
eration.
The heterogeneity
greatly
from that
and
forests
all
stages of degradation and early regen-
this
these habitats differs
of
mature lowland dipterocarp
of
is likely
general orangutan
to affect
Large woody climbers are frequent

forests of the
Lower Kinabatangan
in
the disturbed
floodptain.
behavior and forest use.
Orangutans were studied

at Sul<au.
in this
where they were found
active period
each day
of slightly
to
more than
Based on more than
(612 minutes].
environment
have an average
5 000
hours
hours
of
flowers. Fruits are the

ing
mutually exclusive daily
35 percent feeding;
distance traveled
travel
47 percent resting;
percent moving;
11
percent
daily
was about 300 m. Unflanged
more than other orangutans. When
to the
movement
distance they travel daily
of adult
is
females, so the
significantly less than at
other times.'
Orangutans
seen
at
to eat a total of
Sukau study
the
310 species
site
of plants,
due
that are very
Woody climbers
forests.
their
leaves.
individuals
were consumed
Among
away from areas
19
constant
fruit
supply,
due
to
a rich
to the
year-round
common
lot of fruit
(which
makes up 60 percent
and appear healthy. Compared

primary
forest, they feed
more. Due
to
availability of their leaves
scarcity
Our preliminary
utans possess an
results suggest that orang-
unexpectedly high degree of
behavioral and dietary
them
to
survive
in
flexibility
that
highly logged
males are forced
in
understand
degraded
habitat.
fully
in
and
is
orangutan socioecology
This will be of crucial im-
the design of a sound
management
in
the
Lower
Kinabatangan and other similar areas.
Marc Ancrenaz,
Isabelle
Lackman-Ancrenaz,
and Ahbam Abutani
their
are lighter, they can
orangutans showed
still
move
arboreally.
remarkable
These
flexibility
and
adaptation to the levels of disturbance encountered

(see Box 10.2).
of their diet],'
to
would allow
and secondary
from
populations
to the loss of large trees
continuity, however,
and
to
consume
and travel less and
secondary
However, further longer-term observation
the abundance of
Thus, the orangutans
in
needed
pioneer and climber species that have invaded the

site after logging.
Sukau
forests.
of active disturbance.'^"
On the other hand, the area provides
at
bark; they are used extensively at times of fruit
185 for
for
(9
percent).
were
these 310
for their fruits.
97 for their bark, and
of
are of special importance
strategy for the orangutan population
species. 135
in
Ecological role
The importance
of fruit in
orangutan diets indicates
rest
the vital ecological role these animals could play
and canopy
as seed dispersers, possibly affecting patterns of
move more
along the ground. As females and young orangutans
170
woody climbers
species of trees, 87 species of vines, and 13 species

tablel.
(1
Most foods consumed by orangutans
portance
monocotyledons (see
diet
originate from either pioneer tree species or the
66 families and 156 genera; these included 210
of
the
and insects
percent), flowers (2 percent),
in
unflanged males adapt their move-
consortship,
ments
activities:
and 6 percent unknown. The average
nesting,
males
was spent on these
proportion of time
following
adult females), the
five
food, account-
of
orangutans, followed by leaves (22 percent), barks
direct observation of nine habituated wild orang-
utans (four adult males,
most common
for
forest
regeneration and plant-species diversity."
The species-rich nature
of tropical forests is partly
BORNEAN ORANGUTAN [PONGO PYCMAEUS]
maintained by frugivores moving seeds from the

parent tree to sites
only disperses
some
distance away. Tfiis not
germination sites but can also
sheer size may have enabled
frugivores.^" Their
orangutans
to fulfill
an important role
dispers-
in
large seeds. Larger animals - banteng ISos
ing
increase germination and establishment rates, as
javanicus] and rhinoceros [Dicerorhinus sumatrensis]
by seed-eating animals and seedling
- are virtually extinct on Borneo,'" and the identity
predation
competition are both often higher under the parent
of the
tree.""' The main factor
if
many
plant species
number
the
is
limiting the density of
reaching suitable growth sites.
more
Little
orangutans
in
^'
role
of
and habits. Given
body size and frugivory, orangutans cer-
their large
have the potential
The mechanism
seed dispersal
of
many
disperse
to
is
seeds.
also relevant,
For this to
bats,'"
swallowed whole by oranglikely to
be
dispersed away from the crown of the parent tree,
(such as those
size,
bats,
fruit
Borneo. '^
^'
Gibbons [Hylobates
sunbears
spp.l,
[Hetarctos
barbels IMegalaimidaeL
spp.l,
tree.'"
happen often there must be many
and their populations are also dwindling
(Viverridael,
that are
fruits
the
the tree
seeds by carrying them away from the parent
important consequences for their
Seeds
on fallen
much smaller
Despite their
in
utans and later defecated are more
rely
fruit in
such as Pteropus vampyrus, can disperse large
as the process that the seeds undergo can have

fate.
not known. Even
dropped by an orangutan).
seed dispersal,'" but much can be
inferred from their obsen/ed diet
tainly
the
is
same species as
orangutan, they are unable to access
canopy and would
known about
is
seeds
of their
seed species they disperse
they do disperse the
civets
matayanus],
mouse deer [Tragutus
and pigeons (Columbidael also disperse
fairly
large seeds by defecation, although they are pro-
and dispersed further, than seeds that are spat out
bably unable to disperse the largest seeds taken
or dropped after the fruit pulp has been removed."
by orangutans Isee Table
Orangutans often carry
much
as
tree, by
fruit
as 200 m, before they spit out
or drop the seeds." This
seeds
to
away from the parent
may be
enough
far
escape unfavorable conditions beneath
" Those seeds
the parent tree crown.''
swallowed may take up
to
that are
several days to
defecated. ^^" Such seeds would typically be
800
for
or, conceivably,
be
moved
km
from the
dispersers
as far as 8-10
10.2I.''
""
" Hornbills
(Bucerotidael regurgitate large seeds,'" and travel

long distances quickly'" so are probably
most
important
seed
dispersers
Primates and hornbills eat

subsets of
of
fruit in
similarly
generally,
distinctly
different
is
evidence
in
Borneo;
with
high-fat
small dietary overlap
consume
the
Borneo.
Cameroon."^ There
hornbills
among
in
fruit
Durio
sp.,
one
of the
while orangutans prefer
favorite orangutan fruits
inhabit
Orangutans also function as seed predators,
Ficus sp., a less favored
Borneo's forests. Several are even more frugivo-
destroying around 30 percent of their dietary
but highly important
parent tree.*^
Ivlany
arils
(seed
coverings],
moist sugary fruit."
potential
seed
rous than orangutans (Table

efficient at dispersing the
0.21,
and may be more
seeds. Of 413 fecal
sam-
species." They
may
in
destroy and disperse seeds of
the forest
llefti,
and
food source.
Lynda Dunke
ples obtained from gibbons [Hylobates muetleri X

H. agilis], for
example, 100 percent contained intact
seeds." Similarly, Galdikas'^ found intact seeds
in
9A percent of 6A Bornean orangutan fecal samples,
although this high rate
may have been
a seasonal
effect.
Seed
size is an important factor influencing
seed dispersal by frugivores. The small seeds

figs, for
example,
may be dispersed
of
by abundant
small birds and insects, but the dispersal of large
seeds relies on animals big enough
to carry or
swallow them. These larger birds and

are the chief targets of
ective loss
human
mammals
hunters; their sel-
from an ecosystem seriously impairs
the dispersal of plants with large-seeded
fruits.'"'^
Orangutans are the largest arboreal frugivores

in
Borneo's forests, and
among
the largest of
all
171
World Atlas
Table 10.2 Bornean fauna capable of dispersing large-seeded
Common name
Category and taxon
fruit^
Degree
of frugivore
lEngllsh/Bahasa'l
4 Loss
of this frugivore
would
tiave a
major impact on seed
Pteropus vampyrus
Hy/otiafes spp.
Paguma. Paradoxurus
seeds
dispersal''
large flying fox
dropped
defecated
[musang]
civet
bearcat [binturung]
defecated
defecated
Bucerotidae
hornbill
regurgitated
Ducula spp.
imperial pigeon
defecated/regurgitated
barbel
defecated/regurgitated
Megalaimidae
3 Loss of
tfiis
frugivore
may
Macaca
impact on seed dispersal'
tiave a significant
Pongo pygmaeus
orangutan (mawas)
defecated/dropped/spat
C-D
dropped/spat
C-D
defecated
defecated
Sumatran rhinoceros [badak]
defecated
banteng
A-C
defecated
destroyed/regurgitated
B-C
defecated''
destroyed"
macaque
spp.
2 Loss of this frugivore
may have
sunbear [beruang]
mouse deer
Tragulus spp.
Dicerorhinus sumatrensis
Bosjavanicus
Muntlacus, Cen/usspp-
deer
Argusianus argus
[peianduk]
[kijang. rusa]
Argus pheasant
Loss of this frugivore would have
little
impact on seed dispersal'
Sus barbatus
Loss
[beruk, kera]
minor impact on seed dispersal'
Helarctos malayanus
likely fate
of large
gibbon Iwak-waW
spp-
Arctictis binturong
Most
of
frugivory''
bearded pig {babi hutan]
of this frugivore will
have no impact on seed dispersal'
Presbytis spp.
langur
NasaUs iarvatus
C-D
destroyed^
A-B
destroyed"
squirrel
dropped/destroyed"
porcupine
destroyed"
proboscis
? Effect of loss of this frugivore on seed dispersal
Hystricidae
not known'
is
Ratufa. Callosciurus spp.
monkey
a Animals that rarely eat fruit are not tabulated

b Taxa are grouped according to Corletfs importance index.
unknown
c
Bahasa
is
effect CI
names from
Malay
is
forms the majority
the
Some seeds may
dialect
of the
sanne species, as
used as the national language
annual
single
same
is
in
coded:
individual
species
broken seeds from the
same
in
may
different
intact
and
species." They are
also very wasteful feeders and drop a
lot of fruit,
often unripe, to the ground. This provides food for
172
with A being the
most important;
Indonesia; a sample
is
given
in this
column
of a
few
common
fruit
A,
occasional;
B.
consistent;
c, fruit is
seasonally dominant;
D, fruit
diet,
orangutan feces can contain
terrestrial
'^',
be dropped, or cached and forgotten
process seeds from the

Viâys;
to
Indonesia and Malaysia,
d Degree of frugivory, also adapted from Corlett.
'0'
This index runs from
also tabulated,
feeders,
such as mouse deer
(7"ragu/usspp.], kijang [Muntiacis muntjac],
deer
ICervus
barbatus], and
unable
to
unicoior],
sambar
pigs
(Sus
porcupines (Hystricidae) that are
access the
terrestrial
bearded
fruit in
animals may
the tree canopy." These

in
turn
disperse the
seeds.'""
In
much
of
Borneo, orangutans are declining
because their forest habitat
many
eliminating
being destroyed,
is
Those
plant and animal species.
fragmented forests that remain, but are too small

to
may change
support orangutans,
because
the
of
loss
dispersers. Without detailed

dispersal,
it
in
composition
orangutans as
of
seed
research on seed
impossible to determine exactly
is
these forests are changing.
how
unlikely
is
It
but
possible that there are plant species that rely solely
on orangutans for dispersal which would face

extinction without them.^
More
likely is a loss of
genetic variability within
plant
species that are
dispersed by orangutans and the development
more clumped - and therefore more
A young Bornean
similar effect has
orangutan whose
other areas following the loss or
progress has been
vulnerable - spatial distribution.
been noted
in
significant decline of
an important disperser.'"' "' A
followed at
areas where
the Kinabatangan
smaller-seeded plants
to
shift
of a
ecologically
in
orangutans are absent may also occur.'^
Orangutan Conservation
Project intensive

Food, and especially
fruit,
study
abundance
limits
site.
most
orangutan populations. They would therefore be expected to compete with other fruit-eating animals
for supplies.
Borneo's most frugivorous animals
monkeys
macaques [Macaca nemestrina. M.
are the gibbons, hornbiUs, squirrels, leaf

IColobinael,
fascicularis],
sunbears,
Orangutans show
little
and
arboreal
response
trees, suggesting they are
fruit
as competitors. '' Competition
animals
rences
is
in
some
alleviated to
their food tolerances
civets.
these animals
to
in
rarely perceived
among
fruit-eating
extent by the diffe-
and preferences. The
orangutans' closest dietary overlap
is
probably with
that of other primates.'"
Several features
of
orangutan behavior give
them advantages over other animals. Their bulk

permits orangutans
to
consume
the largest pro-
portion of the fruit crop, and their flexible feeding
and foraging strategies mean that they can take

advantage
This
of
flexibility
limited
may
must
In
Sabah, for example, gibbons fed daily
liana
ICombretum
future supplies for other animals." While orang-
utans certainly compete with other frugivores for
ripens slowly, so only a few ripe fruit could be found
fruits."'
visit
gibbons plucked only
Orangutans
visited the liana
two consecutive days, removing whole bunches

fruit
and dropping the unripe
consume the unripe
fruit of
fruit.
some
Kinabatangan.
one
in
nigrens]; the fruit of this plant
each day, and on each
along the River
significantly
reduce food availability for other animals.

in
living
allow them to be more wasteful
resource, which
study
the ripest
A proboscis monkey
of local differences in fruit availability
on
of
Orangutans also
species, reducing
limited resources,
it
is
the other animals that
may
suffer the effects of this competition.
Niche separation between primates

ieved by differences
in fruit
Orangutans can tolerate very sour or

sympatric primates
may
is
ach-
and habitat preference.
avoid."
In
utans share this tolerance with the
bitter fruit that
Borneo, orangfive
species of
173
World Atlas
HABITAT REQUIREMENTS
Bornean orangutans have
a patchy distribution,
so the broad areas shown
(Map
may
10.1)
the distribution
in
and
map
not accurately represent their true
occurrence within an area
of
forest.
interrupted expanse of forest, their
space and time according
an un-
In
numbers vary
in
to the availability of food
resources and, possibly, the availability of dietary

minerals."' Often there
an intermediate zone of
is
low density between areas
presence and absence,
of
occupied by mobile single males rather than a resident breeding
Bornean orangutans
population.'"
are most abundant
peat-swamp
in
forest
and
flood
swamps appear to prohabitat and some regions
plains (see Table 10.31. Peat

vide a particularly rich
may support
In
quite an even distribution of animals.'"'
more commonly found
general, orangutans are
close
streams,
to
because
of the
and swamps, probably
rivers,
higher incidence of preferred
fruit
trees there. Rivers also act to limit distribution, as

rivers that are
more than about
m wide and 60 cm
deep cannot be crossed by orangutans."
The main reason

from large areas
'"
that orangutans are absent
Borneo may be past hunting
of
pressure. Alternatively, this absence and the large-
may
scale distributional patchiness

reflection of habitat suitability.
inhospitable to
relatively
simply be a
Bornean
forests are
large-bodied frugivores
Sheena Hynd
and, even
Tabin Wildlife Reserve
eastern Sabah.
in
leaf
monkeys
[Presbytis spp.,
Semnopithecus
spp.,
Trachypithecus cristatus]; however, the diet of the

Leaf
monkeys
and
leaves
is
dominated by these types
(supported
by
their
of fruit
fermentative
digestive system], while orangutans prefer other

fruit types."-
'^
The
soft juicy fruit that
prefer" are shared
more
with the
orangutans
macaques and
gibbons.""
habitat.
achieved primarily through differences

Long-tailed
concentrated
availability.
for
in
in
macaque populations are
riverine
areas, while
pig-tailed
are found in highest numbers in dry,

upland areas." '" Of all the primates, the greatest
is
between gibbons and orangutans.
Gibbons are very selective feeders; they manage

feed
in
to
the presence of orangutans by rising earlier
and reaching the main food supplies before the

larger apes. Gibbons also travel
much
longer dis-
tances each day. Their energy-efficient brachiation

(rapid
movement between
tree branches, achieved
by swinging from handgrip to handgrip!
more food sources
174
thrive
slight
example
in
during
times
change
in
dominant
Sabah not
historically
could
to
make
survive.
it
factors
available to them.'^^
makes
poor
of
it
fruit
forest composition,
favor of dipterocarps (which are
particularly
in
the areas of
Sarawak and
occupied by orangutans],
impossible for a breeding population
The hunter-gatherer Penan peoples
influencing
of ecological
they are absent
distribution;
from areas such as eastern Sabah, where the
abundant
forests lack
utilis,
macaques
dietary overlap
the better parts, orangutans find
in
to
demonstrate a comparable example
Niche separation from the macaque species
may be
difficult
hill
sago palm lEugeissona
Arecaceae). their staple starch.

Rijksen
and Meijaard estimated'^' that 35
percent of a typical peat-swamp forest would be

suitable
as orangutan
conservative.
habitat,
but this
More than 90 percent
catchment area
is
of the
may be
Sebangau
used by orangutans.'"' There are
several types of peat forest (see Table 10.31, with

the
highest
orangutan densities being found
shallow coastal peat

Ruling and
swamps
(e.g.
Gunung Palung National
lower densities
in
the deep peat
Sebangau catchment
at
in
Tanjung
Parks], with
swamps
of the
area, even though orangutan
distribution
is
less patcliy
these latter areas.
and more widespread
hunting; or the separation of Barito Ulu from other
in
suitable habitat by rivers and heath forest."
'^
are
abundant
least
in
fiill
forest on ultrabasic soils,
dipterocarp forest,
and
in
both submontane
in
are
in
mountain habitats, although the

breeding population has not been
orangutans
much as
confirmed,'^
Two
halved, depending on the extent of degradation;
populations
may occur between 700 and
as
known.'"
of logging are not
long-term impacts
In
may be
Gunung Palung, some disturbed peat

higher ape
contained
lowland
forest,'^ but
support very few,
The
forest
areas that have been burnt can
away from
drier forests
Ihill
'''
have few
rivers
(flat
lowland] to 30
country up to 500 ml of these areas have
habitats suitable for orangutans.'"
natural habitat
The
heath forest, with
is
vegetation growing on extremely
least suitable
absence
orangutans,
of
low-stature
its
in
the forests of
Barito Ulu (Central Kalimantan) have been
buted
attri-
the high incidence of dipterocarps;'" the
to;
low incidence
large fruiting
of
of
sightings
in
trees;" historical
300
in
Sabah,'" but there have been no sightings
either location since about 1988,"
were thought
to
be
The Kinabalu
of transient individuals.
were found
of a solitary individual
the Maliau Basin Conservation Area,
Sabah,
at
an elevation
of
about
elevations
central
000 m." Similarly,
National Park]
in
northern interior Sarawak.'"'^
The characteristic
altitude
primarily reflects the distribution

favored by orangutans, as there
above
this
pulped
fruit."
altitude
in
the
The presence
Kinabalu and
at high
Tamabu Range (Pulong Tau
the
in
2002
in
in
wandering individuals have been reported
infertile soils, often
over white sand. The low density of primates, and

virtual
Park
in
sets of sightings suggested that
Mount Kinabalu and Crocker Range National
the
Tracks
any, orangutans.
if
food trees and only 25 percent
percent
presence
primary
than
densities
at
of a
and montane forests. Orangutans persist in secondary forest and selectively logged lowland forest,
but their population density
found
rarely
m, although exceptions are

Palung
increasingly being reported. Gunung
National Parl< hosts relatively large numbers of
elevations above 500
and lowland dipterocarp forest and
sities in alluvial
orangutans
Bornean
Orangutans are found at intermediate den-
in
of
is
limit
of
500
a sharp decline
abundance
orangutans
Crocker Range
the
of
food types
of
at
at
soft-
Mount
higher
Table 10.3 Bornean orangutan densities' supported by different habitats
Danau-Sentarum
Borneo 11999 review!
National Park"^
No. of sites
Density
Habitat type
3.0 12.2-3.5)
Flood plain and peat forest
Sebangau peat-
swamp
forest
Density
Density
Density
3.29-4.09
0.91-4.20
4,09-5,87
4,09-587
Shallow peat forest

Mixed
Low
swamp
1.92-2.42
(deep! forest
0.91-1.15
pole" (deep peat) forest
2.04-2.57
Transitional (low pole/tall interior) forest

Tall interior
Alluvial
2.04-2.57
(deep peat) forest
3.22
2.211.2-3.51
and lowland dipterocarp forest
Secondary and
1.28
0.75 10.5-1.01
selectively logged forest
swamp
(old
Mixed
swamp
(recent disturbance! forest
disturbance) forest
3,20-4,59
3,00
Disturbed lowland forest

(hill
1.80
0.5(0.1-1.1)
and dipterocarp! forest
mean number
low
pole forest'
is
of individuals per
2.60-2.77
2,13
forest
3,20
Disturbed peat forest
a Ttie
3.00-4.59
0,85-1,1
Disturbed peat (shallow) forest
Submontane and montane
0.85-4.20
3,33-4.20
Mixed
Upland
Gunung Palung
National Park"
km
a description used
and/or range
in ttie
region for a forest witti
many small
trees.
Adapted from
Rijl(5en,
H D
Ivleijaard,
E (19991. and later sources (see numbered citationsi
175
World Atlas
Box 10.3 PEATLANDS IN SOUTHEAST ASIA AS
The
HABITAT FOR ORANGUTANS
considered to have low fish-species diversity and
rivers of these
peat-swamp
productivity,'^ but this
About 70 percent
260 000
in
km^
of atl tropical peatlands,
occur
Souttieast Asia lespecially
in
Borneo and Sumatra! and
Guinea IWest Papual.

forests
grow over
ficiently
waterlogged
In
environment: not
to infiibit
iversity
all
relatively
is
demanding
particularly
species of lowland forest trees
low
result,
biod-
peat forests, although
in
plant and animal species are restricted to
'"
these environments.^'' '"
local
or suf-
the decomposition of
can tolerate these conditions. As a
some
New
these areas, lowland rain
soils tfiat are flooded
creates a
This
Indonesian
in
so peat accumulates to various
fallen vegetation,
depths.
some
'"
There
and regional variation within

differences
type,'" with
significant
this vegetation
nutrient availability exerting strong influences on

forest composition
Peat-swamp
peat,
carbon
is
refuge for a
in
of
released
both the trees
biomass
the latter""
into
of
the
Much
of
atmosphere
the
number
bird
to
be peat-swamp
and are not associated with any other
wetland or forest habitat. Recent studies have also

highlighted the role that tropical
play
in
peat-swamp
peat-swamp
that
forest
this
ened, and vulnerable species of mammals.^'-
undervalued as a habitat
accumulates,
it
relatively large populations of
orangutans
remaining habitats for
for rare
of the
this
Borneo
'*
'^'
'"'
most important
endangered primate.
nean orangutans are found
in
peat-swamp
Sebangau,
forests
Ithe
habitats with
deep
Kapuas-
and Katingan-Mendawai catchmentsi or
areas with large expanses of shallow, coastal

forests ITanjung Puting
is
of
much
and Gunung
as 40 percent
Borneo's remaining orangutan population
occupy peat-swamp
and threatened
may
forests.
Field sureeys carried out since 1995 in the
River
Sebangau catchment
in
Central Kalimantan,
of
Borneo and Sumatra and
in
Peninsular Malaysia. The true extent and
thickness of these peat deposits are poorly documented.
Thailand
Peatland
100E
176
in
"
Southeast Asia
Most peatlands occur on the islands
v.
'"
Five of the six largest extant populations of Bor-
Palung National Parks].'" As
species, not all of which are wetland specialists."
'"'
Of particular conservation importance are the
of
ecosystem has been
forests
providing habitats for endangered, threat-
peat-swamp
biodiversity
is
and threatened species
of rare
species are considered
specialists,
burned.'"'
As more information on the
forest
has been demonstrated."" Several Southeast Asian
in
in
peat-swamp
These forests provide one
of the large
North
in
Sebangau
considerable and the importance of this habitat as a
Barito,
Peatlands
the River
in
diversity of avian species in
and also when the surface vegetation and peat are
becoming clear
such as those
catchment. Central Kalimantan.'"' Meanwhile, the
following deforestation, drainage, and development
tropical
habitats,
forests represent a significant
former and the thickness

this
peat-swamp
associated with peat-swamp forests."'
because
many
view has changed as
and structure.''^'"'
part of the terrestrial carbon store,
and the
were long
taxa have been found to be associated with
Selangor, Malaysia'" and
hydrology and
both
in
is
new
forests
130E
Top: Canopy of the
peat-swamp
forest in the
John
0. Rieley
John
Rieley
Sebangau catchment.
Above: Impact on peat-swamp forest of the Proyek Lahan Gambut drainage channel.
Indonesia have
shown
to
it
habitat for orangutans.'"' In
areas
forested
experiencing
logging.'''' In
catchment
in
in
ficient to
its
common
Kalimantan,
large-scale,
15.68 km^l
however,
of the
status alone
will,
it
is
illegal
Sebangau
was designated as
government
most
with
indiscriminate
October 200i, most
national park by the
change
be a very important
of Indonesia.
include plans to increase the area of land under

plantation and arable crops (such as
oil
and vegetables]
in
of
people
perimeter
to
who
It
is
support increases
will
recommended
zone be established around the new park,
This
human
large orangutan population. Urgent action
is
and reduce
Peat-swamp
however, be insuf-
save the peat-swamp forest habitat and
forests are
in
decline through-
out Southeast Asia and steps should be taken to
increase international awareness of their global

the carbon cycle and
importance
change processes, as well as
impact
of the
extraction canals associated with
to control
their negative Impacts.
hydrological integrity of the area by negating the

of the
that a buffer
new
activities
rice,
number
be settled around the park
strongly
required to prevent future fires and to restore the
illegal
palm,
the
in
orangutan and other
in
climate-
for the conservation
wildlife.
logging that hasten the runoff of water from
the peatland ecosystem.
Other threats to the Sebangau National Park
John
Susan
Page,
and Suwido
H.
Limin
0. Rieley,
177
World Atlas
may have been due
altitudes
and durians [Durio
spp.l
is
It
required
difficult to
population
healthy
Borneo claims an extensive
of
of
community
the lands they clear for farming
to
known as kebun
(areas
or 'gardens']; these rights
are inherited. The 'gardens' are maintained
orangutans. They have a complicated and poorly
system
understood social and ranging system
response
when secondary vegetation reclaims

Community land extends beyond the
and food
areas and includes
communities that travel long distances

to
mast seasons, presence
to
be confident
of either the
number
it
loose
of
in
of females,
shortages. With our current knowledge,
is difficult
of individuals
""
or the area these individuals require. '^^
in
followed by a fallow period
of cultivation
swamp and
the
site.
cultivated
virgin forests that
are conserved for the collecting of wild produce and
hunting of animals.^'
Hunting of orangutan was widespread amongst
required to preserve the orangutan social structure,
the Dayak peoples, generally using a blowpipe and
The apes were considered
poison.
be a prized and
to
POPULATION STATUS, TRENDS, THREATS
delicious food item
Human
Human
sexual potency.''"^ Infant orangutans were kept as
attitudes and traditions
settlements have existed
40 000 years.'
least
caves at Niah
historic people,
of
in
were
and clans
Sabah shows
in
who may have caused

still
for at
a regular food item of pre-
orangutans from several regions
tribes
Borneo
Archaeological evidence from
Sarawak and Madai
in
that orangutans
have
of
the extinction
Borneo.
Some
a religious association with
pets
and also a medicine
longhouses or
in
sold.
Under some
from eating orangutan meat;

the species from extinction
this
in
of the flood plain
Kapuas River and the
of the
Mentarang area
in
member
of a particular species Ifual and,
comes
form, can help his or her
in this
Kayan
is
some
inhabitants
Borneo and the more
known
collectively
as Dayaks'. These Austronesian peoples
came
to
Borneo from the Malay Peninsula and Sumatra
waves
starting
some 4 000
it
in
living
years ago. This
was
descendants.
tribes today;
one study
among seven
the Batang Ai region of Sarawak,"
in
was permissible
hunt the apes
to
to obtain
head on special occasions.'"
Orangutans were also hunted
unclear between the ori-
ginal prehistoric inhabitants of
in
households
though
East Kalimantan.'"
The relationship
recently arrived
valleys of the
area
of areas."
that after death an ancestor be-
found the orangutan to be a tua species
much
traditional
The Iban believed
are absent from the largest part of Sarawak's low-
Brunei, and from
ensure
may have spared
number
This belief continues
land forest, from north of the Rajang River, through
to
systems, certain families or clans were restricted
the apes, often involving hunting them. Orangutans
all of
some Dayak
reasons. To
a soul'"
and
for religious
peoples, they possessed
this 'soul substance'
was considered
Some
higher than that of humans.
Iban groups
regarded the orangutan as being a representative

of their
war
custom
of
god.' Related to this belief
the
Kelabit and
was
the
Kayan peoples that
part of the great island-hopping expansion of the
involved rubbing dirt and ape hair into the breast of
Austronesians from their original base
On Borneo, the Dayaks

ties
adapted
interior, the
to
meet
diversified to
in
Taiwan.''
form socie-
a variety of conditions. In the
dominant form
of land
use was shifting
cultivation, originally of root crops, with
hill
being adopted around the 5th century.
Hunter-
arose
in
some
areas. The Iban,
populous people
in
parts of
now among
the most
Sarawak and West
Kalimantan, are relatively recent arrivals from
Sumatra. They invaded Borneo along the Kapuas

River
in
West Kalimantan during the Uth and 15th
centuries. Resident Dayak groups
were absorbed or
among
traditional
systems
used
prevent
child, to
The skin and
until the
it
being stolen by an
hair of orangutans have
been
present for war cloaks, jackets, and
caps, and to decorate the handles of swords.'"
Compared
ture
only
to
other animals, orangutans fea-
irregularly
in
traditional
mythology.
Linnaeus related travelers' reports that suggested
human rulers, who had fallen from

some Malay people, orangutans are humans who refuse to speak for fear of
being enslaved.'" The Iban seemed to view orangutans as their forest cousins; some stories suggest
they made no clear distinction between humans and
they were once
grace." According to
apes." Indeed, traditionally prepared trophies
displaced as the Iban spread.
Under
newborn
ape.'^'
rice
gatherer societies, such as the nomadic Penan,
178
tract
Within these territories, individual families
land.
have rights
spp.l.'"
quantify the continuous area
support
to
den-
to unusually high
such as mangosteens [Garcinia
sities of fruit trees
of
the Dayaks, every longhouse
land tenure
community
in
human
of
skulls have been found mixed with orang-
utan skulls.'"
Many
stories tell of attacks,
and
BORNEAN ORANGUTAN [PONCO PYCMAEUS]
rapes,
women by male orangutans, and

men" by female orangutans. In
stories, women were often kept as the
of
(occasionallyl of
these
orangutan's lover or bride;
in
some accounts
inter-
as traditional Dayak
increased
erged
strength of orangutans
every
pythons,
with
fights
in
The Kenyah and Kayan peoples
are unable to look straight into the eyes of an
orangutan
case they offend
in
hunt and eat
but they will
it,
still
are
in
in
1997 when an economic
Indonesia.'
week two
infant
smuggled
out
Products such as hornbill
of
and rhinoceros
'ivory'
Borneo have been traded
of
interior of
international trade
Borneo
Major movements
of
Growth
the
in
early
of
19th
massive cultural changes.

indigenous peoples
of
were induced by the head-hunting habits
Borneo
of the Iban
around the same time. The constant threat

warfare and transition being undergone by
of
many
societies diluted the traditional mythical status of
orangutan.''^ This
into
killed to obtain
The current position

The Bornean orangutan
by
it
has a very high
classified as
Endangered
indicating
risk of extinction in the wild in the
near future." By 2004, the
remaining area of
total
Bornean orangutan habitat was around 86 000 km^

supporting 45 000-69 000 individuals. Their
bution
IS
distri-
summarized below."'"
Central Kalimantan has several forest areas
of
large
enough
to
vantage
individuals.
system and
regulatory land-use
religious constraints on hunting the

of traditional beliefs
ape vanished.
wurmbii\:
thought
to hold over
The peat swamps
of the
6 000
32 000
western
and the
nearly 2 500.
Young orangutans
at
an
orphan sanctuary
in
Borneo.
Commitante
selling previously
timber companies or
illegal
were superseded
and
more
Coupled with these changes was the
arrival
advanced
technologically
p.
support viable orangutan
Gunung Palung National Park

Raffaetla
loggers. Old logging techniques
is
IP.
Sebangau catchment may hold
and community respon-
removed constraints on
to
is
populations, and
untouched forest
that
Central Borneo orangutans
who came spontaneously to take adnew trading and farming opportunities.
traditional
mothers
these infants, or the infants
from the pet trade.
Indonesia, or
more
'^^
who came
as transmigrants sponsored by the government
by
exist.
Borneo by
Javanese, Maduranese, and other people
Loss
to
process was accelerated
more recent mass migrations
sibility
Batam
via
way deep
their
return.
in
relations
century precipitated
The
Jakarta
of
that die before being confiscated, the
that are not rescued
and bronze gongs, have found
of
orangutans from Kalimantan
course, take into account those infant orangutans
more." Trade goods from China, such as ceramic
by
em-
percent of the estimated total
were
the
crisis
has been estimated that
young orangutans confiscated by
as far afield as China for a thousand years or
into the
of
population at that time." This figure does not, of
horn from the interior
jars
It
Borneo and several other trade routes also
2000 represents
Recent history
may be
Singapore."' Orangutans are poached throughout
The number
it."
although they
and the associated pet trade are sizeable; they
ing
breeding occurs. Other stories describe the great
crocodiles, or bears.
relics,
recent provenance. Orangutan losses due to hunt-
destructive methods."
of
European people seeking
'scientific collections'
One
or opportunities for sport hunting.
shot 217 orangutans
in
one small
around AO individuals were
The spread
of
common
that
many people no
orangutans as potential food items;

(forbidden)
in
the 1890s.
the Muslim culture into
Borneo has meant
under Islamic
army personnel
biologist
area;'^' 'bags' of
much
this
is
law. Nonetheless,
Inot all of
whom
either Indonesia or Malaysia) have
of
longer see
haram
some
are Muslim
been reported
in
to
go on poaching expeditions.
Some
foreign tourists
still
purchase ape skulls
179
World Atlas
Borneo
Northwest
orangutans
IP.
(also a
Ramsar
forests
in
and the surrounding peat
West Kalimantan are estimated
about
hold
Site)
1997-1998
'^'
There are
orangutans remaining
last viable
in
population of
Batang
the
to
500 individuals following the
fires.
few
Sarawak, with the
UO
relatively
to
760 being
in
National Park and adjacent
Ai
Lanjak-Entimau Wildlife Sanctuary."

Northeast Borneo orangutans
Sabah
18
is
thought to
000-18 0001
3 000
in
hold
p.
mono]:
around
11000
individuals.' with
IP.
A Population and Habitat
p.
pygmaeus]: Danau-Sentarum National Park
about another
workshop held
estimated
to
of
from
lation
this
be less than
at the start of the
H percent of what
Holocene,
000 years ago,
is
was
when
it
merit conservation attention.
decline of the species accelerated towards the end

of
the 20th century, with deforestation and degra-
dation of Bornean forests since the 1970s.

unlikely event that all
human
In
the
disturbance and
Feeding time at the
anthropogenic mortality suddenly ceased, orang-
Sepilok rehabilitation
utan
center
in
Sabah.
numbers would recover
per year due
to their
at only 0.006
percent
very low reproductive rate."
that the
would seriously
The workshop's selection procedure con-
sidered
taxonomy, habitat diversity (uniqueness
and
and distribution among major
peripherality],
and provinces).
political units Istates
(73 371 km') occupies about 10 percent
Borneo; about half remains under
natural forests," although
palm and pulpwood
oil
plantations continue to expand. Eastern Sabah
almost completely uninhabited
lowland forest,
The
fails to
unit."'"
now
10.4).
it
jeopardize the taxon's integrity as an evolutionary
became
from Sumatra (Table
popu-
of a
was noted
It
"loss of any of these populations
the last ice age ended, sea levels rose, and Borneo
isolated
Bornean
does not imply that
list
the
to offer
the
orangutan ITable 10.5);'" the absence
Sabah
Bornean orangutans today
were considered
10 habitat units that
East Kalimantan.
number
total
Assessment
identified a set of
greatest potential for sustaining
of the island of
The
Viability
January 200A
in
late
until
was
about 1960, but
only 25 percent of land area remains under
1960s
managed
in
much
ways
of the state's
is
it
heavily logged.
From the
that resulted in severe depletion
now
timber reserves,'" and they are
virtually exhausted.""
land area
of
the 1990s, Sabah's forests were
into
Close to 8 percent of Sabah's
included
the state's system
in
of
protected areas, but about 60 percent of the orang-
utans
Sabah
in
live
outside protected areas,
in
production forests that have been through several

Elaine Marshall
rounds
of
timber extraction and that are
still
being
exploited for timber^
Sarawak (124 500 km')

state and, like Sabah,
area
Malaysia's largest
land
of its
included or proposed for inclusion within
is
state-protected areas.
of
is
around 8 percent
It
has been
major exporter
timber since the 1960s; log production increased
steadily during the 197Ds
and 1980s
more than
to
U million m^ per year More than 2 000 km' of forest

were logged each year during the 1990s, a process
that continues to date.
of
Much
orangutans due
bited by
forest
of
Sarawak
is
uninha-
conversion
to past hunting,
by shifting cultivation, or for ecological
reasons. Significant populations of orangutans
occur only
in
the
around the Batang
Entimau
Wildlife
Betung
with
south-central interior,
Ai
to
now
and
National Park and Lanjak-
Sanctuary, which
Kerihun
National
Kalimantan. These protected areas
thought
in
be subject
to
is
contiguous
Park
in
relatively
in
West
Sarawak are
little
threat,
although they are understaffed and vulnerable to

illegal
logging
and hunting, including probable
cross-border logging from Kalimantan.'"
The four provinces
180
of
Indonesian Kalimantan
BORNEAN ORANGUTAN [PONGO PYGMAEUs]
1536 000
of
km'
occupy more than 72 percent
in total)
Borneo; orangutans
Threats
to
still
occur
Table 10.4 Bornean orangutan population decline since 8000
three of these.
in
abundant and widespread, however, and have been
Year
Estimated number of
Decline from
Bornean orangutans
previous estimate
many years. An example is the

Proyek Lahan Gambut million hectare mega-
8000 BC
420 000
reclamation program
Central Kalimantan. This
1900
230 000
supported by former President Suharto,
200A
up for
building
project,
in
was designed
to boost the faltering transmigration
program and
rice
was responsible
swamp,
utan
production
the mid-1990s.
in
km'
15 000
for draining
including up to 7 000
km'
of
and the land was burned
appears on
satellite
deforested
land.
images as
45 percent
57 000 145 000-69 0001

Base d on Singleton,
1..
e(a(. 120041
75
and Rijksen, H.D
l5l
percent
Meijaard, E I1W9I
It
peat
of
prime orang-
The project was abandoned
habitat."^
1998,''
BC
populations and habitats are
their
2002;
in
it
Table 10.5 Bornean orangutan populations critical to species integrity
in
now
a circular patch of
Estimated population
Habitat unit
Subspecies
Sebangau
300
P.
additional cause of forest loss; by 1999, slash-and-
Tanjung Puting
000
P. p.
wurmbii
burn agriculture was reported
Belantikan
000+
P. p.
wurmbii
Mawas
3 500
P. p.
wurmbii
Gunung Palung
500
P. p.
wurmbii
Sabah Foundation
320
Subsistence agriculture
is
an
have affected 27
to
percent of land area
in
Kalimantan, 87 percent of
which was presumed
to
have been prime orangutan
habitat at
some
stage.
and
is
palm and
been an issue
Acacia.'^ Coal mining has long
Kutai National Park,
oil
becoming one
in
in
other
Kalimantan
tires
damage occurred
province.
East
early
the worst-
in
Kalimantan." Localized
in
percent
was lost.'^ The dispeat-swamp and lowland
forest
was
Kutai National Park. 95
lowland forest
proportionate
loss
of
the most serious consequence as this
the richest habitat for orangutans.'^ Large

of
orangutans were
the flames and
Shockwave
forests,'"
in
'refugee crowding'
fires,
Borneo's orangby 33 per-
fires also
in
occurred
Borneo's cities."""
The remaining Indonesian forests

officially
to
adjacent
in
may have been reduced
one year'" Serious
health
11
5001
bution range of the orangutan

it
is
to
that are
timber production, watershed protection, or bioOfficial
pygmaeus
P. p-
pygmaeus
1.,
et
a(.
120041.
in
Kalimantan, and
generally thought that orangutans are unable
term
survive long
heavily logged
in
forests,'
although the northeast Borneo orangutan appears

be exceptionally resilient
is
It
that
after
the
illegal,
to logging (see
Box
to
10.2).
rather than the legal, activity
causing the current deforestation crisis
is
timber concessions
almost completely overlap the fragmented
distri-
repeated
national parks
decline
in
logging.
is
in
logging
in
the
huge
to
stop
these areas will inevitably lead to a
further dramatic reduction
orangutan
Illegal
rampant and has caused
orangutan numbers. Failure
illegal logging in
population
in
forest cover
size.'"
and
in
Danau-Sentarum
National Park and Betung Kerihun National Park,

tor
intended to be permanent are dedicated
diversity conservation.
mono
mono
P. p.
Based on Singleton,
verted to unplanned, private plantations or burnt
200^, with the resultant haze again threatening
human
500
smoke during and after the fires;

may have precipitated
causing stress that reduces breeding
in just
I'l
P. p.
Kalimantan, however. Vast areas are being con-
of
utan population
> 2 500
P. p.
people while fleeing
killed by
success.' As a result of the
cent
Batang Ai/Lanjak- Entimau/
is
numbers
the displacement of apes

a
I'')
1980s.
damage was extreme;

of
3 000
during 1997-1998,
burning around 52 000 km' of forest

affected
4 000
and droughts have ravaged
repeatedly since the
Significant fire
Kinabatangan
Gunung Gajah/Be rau/Kutai
Danau-Sentarum land surrounds
sought inside additional national parks.''
Furthermore,
mono
Betung-Kenhun
areas, such as Barito Ulu, with mining rights also
being
P. p.
lorestry concession least]
Indonesian forests are also being increasingly
converted to plantations, especially for
p wurmbii
example, are being destroyed by
illegal logging
and have almost no capacity or support

this;
to prevent
Tanjung Puting and Gunung Palung National
Parks have also been badly affected. The construction of canals for illegal logging drains
destroys deep
peat-swamp
forests,
and
and represents
181
World Atlas
major threat
orangutans (see Box
to
Current trends suggest that

forest will be lost or badly
the
mam
forecast
correct,
orangutans
in
If
Kalimantan, and
the
this
Kalimantan.

field
the
late
'^^
National Parl<
Kutai
1960s, at
and
region of Sabah, Malaysia."''
began
tigation
in
1971
A long-term
"'
now designated Tanjung
the area
Park,""":'" and continues
reserve
modern technology,
monitored by
is
habitat
satellite,
in
sup-
teams may be
slow-breeding
Slow-moving,
inves-
which depend largely on
in
Bornean
Puling National
Danum
studies have included work at
of
dispatched to counter them.'''
fruit in
orangutans,
fruit-poor lowland
forests, are very vulnerable to rapid low-
land forest clearance, whether for plantations or
present day. Later
to the
to protect
forest inhabited by orangutans. In an
land uses and wildfires so that
East
in
Central Kalimantan,
in
proposals
which comprises 5 000 km''
This remote sensing captures evidence of illegal
in
Kinabatangan
Ulu
the
in
Mawas
in
in
other parts of
in
plemented when necessary by aerial photography.
study of Bornean orangutans began
Kalimantan.'^'
involved
is
area,
exciting application of
the
The
Mawas
peat-swamp
hard to see a future for wild
is
it
the Balikpapan area as well as
degraded by 2010, with
driver being timber extraction."
is
Foundation rehabilitates and releases orphans
10.3].'^
Indonesian lowland
all
through forest
Valley, the
incidental to
fires
logging and
Kinabatangan floodplain, Tanjung Puling National
drought. The destruction of forest ecosystems on
Park, Kutai National Park and elsewhere, on feed-
the island of Borneo has an historical
and beha-
ing behavior, adaptation to disturbance,
vioral adaptations of rehabilitated individuals.
The
chief experience of researchers
1970s has been
since the
damage and
of
tected areas;
Borneo
in
fires.
As
may
well prove
utan, at about 57 000,
more
decline.
in
conservation, often through non-
governmental organizations. For example,
is
It
symbolic
island. Potentially viable
project along the Kinabatangan River, supported by
forests
WWF-Malaysia and working
only focused investment
Department.
Wildlife
was
involved
the
1980s
in
statewide conservation strategy and

field
studies to support
National
Sabah
Sarawak, WWF-Malaysia
In
during
management
Batang
to
more
Indonesian Borneo, the Orangutan Found-
beginning to
forest restoration.
Nature Reserve,
Strict
into both conservation
improve land use and local gover-
and
The Borneo Orangutan Survival
in
some areas
are beginning to work
and that cross-border cooperation
forests,
National Park, rehabilitates and releases orphan
Lamandau
and sustained
the Bornean states and
closely with local peoples to safeguard their
ation International funds patrols in Tanjung Puting
in
addition, coordinated
all
will require not
be cleared and burned. There are signs that local
governments
and supports research
them
and around the areas
in
nance, and to resist the factors that dispose forests
Park and the Lanjak-Entimau Wildlife
orangutans
In
territories to
Ai
Sanctuary.
In
in
damage
orangutan populations and
but to secure
be required by
effort will
conducting
of
still exist,
themselves.
preparing a
in
the Bornean orang-
of increasing habitat
Hutan runs a community conservation and research
closely with the
of
a snapshot of a species
is
across the world's most biologically rich tropical
Sabah,
in
but
difficult to protect.
The current population
a result,
most researchers have either moved on or become

involved
cannot
become complete,
will also
and tire-maintained scrub, grass, and farmland
populations, with
and forest
it
most pro-
We
moist forests surrounded by flammable
isolated
orangutans coming under increasing pressure from

logging, land clearance,
largely irreversible.
is
it
whether
predict
escalating habitat
declining wildlife
momentum
that will be difficult to restrain even in
become
is
easier both bilaterally and as
facilitated
by the Association of Southeast Asian
Nations.
can only be hoped that these efforts
It
will
continue and multiply.
FURTHER READING
Ancrenaz, M., Calaque,
R.,
Lackman-Ancrenaz,
I.
(2004) Orang-utan nesting behavior
Malaysia: implications for nest census. International Journal of Phmatology 25
Ancrenaz. M.. Gimenez,
0..
Ambu.
12005) Aerial surveys
give
L.,
Ancrenaz.
K..
new estimates
Andau.
for
P..
Goossens.
orang-utans
in
B..
Payne.
Sabah,
http://dx.doi.org/10.1371/journal.pbio.0030003.AccessedDecember8 200i.
182
151:
J..
in
disturbed forest of Sabah,
983-1000.
Tuuga.
Malaysia.
A..
Lackman-Ancrenaz,
PLoS Biology 3
11):
I.
e3.
BORNEAN ORANGUTAN IPONGO PYCMAEUS]
Bennett, E.L. (19981 The Natural History of Orang-utan. Natural History Publications, Kota Kinabalu, Sabah.
Currey,
Doherty,
D.,
Indonesia.
Goossens,
Lawson,
F.,
SE Asia and
S.,
Newman,
International
B., Chikhi, L., JaliL, M.,
J,,
Ruwindrijarto, A. 120011 Timber Trafficking. Illegal Logging in
Consumption of
Illegally
Sourced Timber. EIA and Tekepak Indonesia.
Ancrenaz, M., Lackman-Ancrenaz,
Patterns of genetic diversity and migration
pygmaeus] populations from Sabah, Malaysia. Iôlecular Ecology

Jepson,
MacKinnon,
Jarvie, J.K.,
P.,
K.,
Mohamed,
I.,
M.,
Andau,
P.,
Bruford, M. 120051
increasingly fragmented and declining orang-utan IPongo
in
Monk, K.A. 120011 The end
(21:
Wl-456.
for Indonesia's lowland forests''
Science 292:
859-861.
Maltby,
Immirzi, C.P., Safford, R.J., eds (19961 Tropical Lowland Peatlands of Southeast Asia. lUCN, Gland,
E.,
Switzerland.
Morrogh-Bernard,
pygmaeus]
Husson,
H.,
S.,
Page, S.E., Rieley, J.0. 120031 Population status of the Bornean orang-utan [Pongo
Sebangau peat swamp
the
in
forest. Central
Kalimantan, Indonesia. Biological Conservation 110 HI:
U1-152.
Rieley. J.O.,
Page,
eds (1997) Biodiversity and Sustainability of Tropical Peatlands. Proceedings of the
S.E.,
Symposium on
International
Biodiversity,
Environmental Importance and Sustainability of Tropical Peat and
Peatlands, Palangka Raya, Central Kalimantan, i-8

Rljksen, H.D., Meijaard, E. 119991
Our Vanishing
September
Samara
1995.
Relative: The Status of Wild
Publishing, Cardigan, UK.
Orang-utans
at the Close of the Twentieth
Century. Kluwer Academic Publisfiers, Dordrecht.
Ross, M.L. 120011 Timber
Booms and
Institutional
Breakdown
in
Southeast Asia. Cambridge University Press,
Cambridge, UK.
Russon, A.E. (1998) The nature and evolution
orangutans [Pongo pygmaeus]. Primates 39
of intelligence in
lAl:
i85-503.
Schwartz,
Wright,
J.,
S.J.,
famine
ed. (19881
Orangutan Biology. Oxford University Press, Oxford.
Carrasco, C, Calderon.
in
0.,
et aL (19991
The
El Nifio
southern oscillation, variable
production, and
fruit
a tropical forest. Ecology 80: 1632-1647.
MAP SOURCES
Map
10.1
Orangutan data are based on the following sources:
I.
(20041
Orang-utan Status
in
Sabah: Distribution and Population
Size.
Kinabatangan Orang-utan Conservation Project, Sandakan, Malaysia.

Meijaard,
E.,
Husson,
Lacy,
S.,
et ai (forthcoming) Orangutans: out for the count or hanging on''
R.,
lin
preparation
for Science].
Meijaard,
E.,
Dennis,
Meijaard
(2004) Borneo Orangutan
R.,
Singleton,
\.
& Dennis
120031 and
amended
PHVA
Habitat Units: Composite dataset developed by
by delegates at the Orangutan
PHVA Workshop,
Jakarta, January 15-18
2004.
Singleton,
R.,
I.,
Wich,
Byers,
S.,
0.,
Husson,
S.,
Stephens,
S.,
Utami Atmoko,
S.,
Leighton. M., Rosen, N., Traylor-Holzer,
eds (2004) Orangutan Population and Habitat
Viability
K.,
Lacy,
Assessment: Final Report. lUCN/SSC
Conservation Breeding Specialist Group, Apple Valley, Minnesota.

ACKNOWLEDGMENTS
Many thanks
Husson
to
Mark Attwater (Orangutan Foundation),
(University of Palangkarayal, Ashley
Raffaella
Commitante
(University of Cambridge],
Leiman (Orangutan Foundation], Helen Morrogh-Bernard
Palangkaraya], and Rondang Siregar (University of Cambridge] for their valuable
comments on
Simon
(University of
the draft of this
chapter
AUTHORS
Kim McConkey, UNEP
Box
W/orld Conservation Monitoring Centre
10.1 Julian Caldecott,
UNEP World
Box 10.2 Marc Ancrenaz. Isabelle Lackman-Ancrenaz, and

Box 10.3 John 0.
Rieley, University of
Ahbam
Nottingham, Susan
E.
Abulani, Hutan Foundation
Page, University of Leicester, and Suwido H. Limin,
University of Palangkaraya
183
World Atlas
Anup Shah/naturepl.com
184
SUMATRAN ORANGUTAN [PONGO ABELIl]
Chapter
Sumatran orangutan
[Pongo a be Hi]
McConkey
Kim
orangutans [Pongo abelii Lesson,
Sumatran
are
18271
now mainly
restricted
11-13
to
isolated forest units, found mostly to the north

of
Lake Toba on the Indonesian island
of
Sumatra
(lâp
11.11" The absence of orangutans farther
south
is
considered to be the result of hunting over
many hundreds (perhaps even thousands]

although there are records
of
of years,
Most are found
lAcehl.
1
now
the province of Nanggroe Aceh Darussalam
lowland areas, below
in
000 m, although transient individuals - usually
males - are sometimes seen
The most
at
in
is
set! of
north-
the Leuser Ecosystem (the
large landscape within which the
National Park
in
Gunung Leuser
Aceh, and particularly
in
large
possess a mental
aware
the
They are able
area.
map
seasons. There
of fruiting
of fruit.
Orangutans observe the
and have been seen

Fruit
Valley
those seen
sities of
to
in
and the Kapi Plateau. The

in
soil
these areas are similar to
the lowlands, allowing orangutan den-
occur^'
Two
isolated
in
and
their patchy distribution within broad
of forest at
lower altitudes are both thought
Orangutans consume a variety
of fruit types,
most months. Over three years
Gunung Leuser National

observations were of
at the
same time
to a tree
Park, 58 percent
fruit,
taken from
may be
bearing
of
fruit
attracted
with soft
such as Antiaris toxicaria (Moraceae],
Cyathocalyx sumatranus (Annonaceae],
Mallotus
Carcinia spp.
(Clusiaceae], the
in
forest to
is
in
the
Lumut
coastal
not considered viable.'"
schaeorocarpus
rambutan Nephelium lappaceum,

"
and Xerospermum spp. (both Sapindaceae]."

Large
fruits with
volume greater than
such as durians [Durio
71 cm^,
Bombacaceae] and
spp.,
jackfruit [Artocarpus elastica, Moraceae], are also
favored, although smaller fruits are usually
In
and feeding behavior
the Suaq Balimbing
swamp
in
more
the diet.""
forests of
Gunung
Leuser National Park, the following produce almost
Sumatran orangutans have

and appear
in
Ketambe
at the
abundant and feature more regularly
fruiting trees
which
92 different tree and liana species." Several
juicy pulp,
be
phytocrenoides (Menispermaceae],
also a very few orangutans
Diet
to
availability of soft-pulped fruits.'
together form the largest portion of their diet
site in
the Padang Sidempuan,
known as East SaruUa. There are
a population that
000
expanses
Batang Toru forest block], and the other
swamps,
paths."
Tinomiscium
Tarutung, and Sibolga area (known as the West
the east of there,
flights of hornbills
(Euphorbiaceae],
three to six individuals per square kilometer
occur farther south, one
also evidence
typical restriction to altitudes of less than
these usually semisolitary apes
Mamas
to
be
the behavior and distribution of orangutans. Their
also sustain orangutan populations, these being the
upper
to
has a pivotal influence on
availability
of feeding
type and ecosystems
is
to follow their flight
swamps and certain lowland parts of the Alas

Valley." Two highland areas in the Alas Valley area
coastal
and
of food sources,
to
that they read various signs indicating the presence
governed by the
higher altitudes.
viable populations are found
ern Sumatra, including
of
recognize important food species, are thought
orangutans farther
south as recently as the 1830s." The majority

live in
topography
locating
continuous supplies
be familiar with the
(Tetrameristaceae],
little difficulty
to
of fruit:
Tetramerista glabra
Sandoricum
beccarianum
185
World Atlas
Map
186
11.1
Sumatran orangutan
distribution
this
chapter
IMeliaceael, and Neesia
glabra IBombacaceael.
cf.
These are considered staple orangutan foods
In
Although there
is
evidence for seasonal dietary
Sumatran orangutans, some
in
almost continuous availability
monthly variation
example,
in
diet
Moraceae)
so
cf at least eight
strangler species were available for eight months of
the year
in
the apes
may move
'^
Ketambe.''"
to
around
orangutans.^" Strangler
densities
the
recorded as
fruit
eaten by
of
all
fig
trees occur at very high

large crops of
easily digested fruit at relatively short intervals."
Certain
species are favored over others, and at
fig
Ketambe they include Ficus annulata,

F.
"
drupacea,
F.
stupenda, and
Seeds are
often rich
F.
in fat
F.
benjamina,
the
Sumatran orangutans, and
in
as opportunistic rather than as the
interpreted
outcomes
planned hunting expeditions." " No
of
vertebrate remains were found
in
feces during a
Ketambe; individual orangutans
in
eggs and sometimes also
inspected squirrel nests, so
may
eat nestlings
observed eating an infant gibbon iHylobates

slow
[Nycticebus
loris
lorises
coucang].''''
were also seen
if
An adolescent female was
the opportunity arises."
some areas and produce
in
rare
is
did occasionally eat bird
have been
figs
Carnivory
lin
those incidents that have been reported have been
supplies, but there
'
consumed,
stomach.
of kaolin-rich clay) to settle the
new
is
also
is
provide mineral nutrients or
to
three year study
to find
half
least 17 species of insects are repre-
the orangutan diet. Soil
does become scarce
switch to less-preferred foods,""
Ketambe,
At
in
fruit
If
such as bark and leaves."
providing
case
sites offer
of high-quality fruit
accordingly slight." For
is
figs [Ficus spp.,
also a tendency
sented
presumably
these habitats.""
changes
" At
eaten.'"
lar]
or
and seven slow
be eaten by three adult
to
females over 20 years
Ketambe and Suaq
at
Balimbing." Lorises are easy, slow-moving prey

(although with sharp teeth], and orangutans are
equipped
ill
catch faster-moving animals.
to
subutata.'
with a high calorie
Ranging behavior
content, to provide the seedling with early suste-
nance after germination. As these same attributes
orangutans with quite diverse ranging patterns. As
make seeds
noted
attractive to animals, in tropical forests
seeds are often protected by hard cases or

strong
Their
hairs.
teeth
irritant
and jaws," manual
and strong hands, allow orangutans
dexterity,
to
overcome these defenses and obtain the seeds

of various species,
Heritiera
elata
including the favored fruit of
nutritious
belowl.
seeds
The
of
9,
in
cf.
malayana^'-
"
Isee
these seeds, combined
in
the
Leaves are also regularly eaten by Sumatran

orangutans, representing 5-25 percent of feeding
The apes usually eat new shoots
consume
or buds, but also
selected species.
and 'wanderer.'" This conclusion was based on data
from Ketambe, where 'commuters' were said
make up 60 percent
live
home ranges."
to
the mature leaves of
high tolerance of generally
Dendrocnide spp. lUrticaceael, although
they take great care to prevent their lips coming

contact with the leaf surface." Other items
consumed include
aerial
stems
of
roots,
epiphytic
fungi,
climbers, grass, and leaf
orangutans also use their strong teeth and
to strip
bark from trees. Even the phloem and
xylem layers
of
in
At Suaq Balimbing, up
15 subadult males have been seen to use a forest
square
the
wood
of
certain
trees are
200
of a
grid of survey paths. Clusters of
home range boundaries

Some subadult males
related females sharing
have also been
may range
identified.
very widely, but there
presence
for the
lifetime
jaws
to
with
16 adult females, nine adult males, and at least
may be
galls;
population,
but stable and widely
large
individuals
unpalatable species has been demonstrated. For
orchids, the
the
of
overlapping
example, orangutans eat the leaves of stinging
into
classes
of three social
patch of only h ha, the area bounded by a single
forest.'""
nettles,
terms
linked to ranging behavior: 'resident,' 'commuter,'
Data from Suaq Balimbing indicate that most
Neesia
orangutans an advantage over other primates
'
these have been interpreted by
Chapter
some observers
in
be used by several
to
30 percent residents,' and 10 percent wanderers'.
ability to exploit
observations."
likely
is
at
Orangutans
with a tolerance of unripe and acidic fruits, gives
same
area
tools to extract the highly
|Sterculiaceae|.'
Suaq Balimbing also use
single
was no evidence
commuters. Ranging behavior

terms
of very
large
ranges, parts of which are used
more
better described
home
persistently than
social
of
in
depending on various
others,
and ecological
factors. Differences
in
resource driven, as the Suaq Balimbing

forests offer
more
between
ranging behavior
plentiful
may be
swamp
and continuous supplies
of nutritious fruit.
The home ranges
of
male orangutans are
consistently at least two to three times larger than
187
World Atlas
km'
25
swamp
the
in
and 8 km'
forest
in
Ketambe.^" The Suaq Balimbing orangutans may

be using areas
make
of
swamp and
use
best
of
the
hill
forest
in
order to
and temporal
spatial
patchiness of food resources." The greater range

overlap seen
in
swamp
the
forest probably results
from orangutans sharing access
to
areas with a
high density of favored food trees."
movements do occur regularly

when trees from the family
Food-related
during mast fruiting,
Dipterocarpaceae flower
synchrony,
in
usually
along with several other species, to provide a
superabundance
mast period
normal
of food. In
the lowlands tend to be
more
1997
in
fruiting
seasons
productive, but during
some orangutans
in
the
Gunung Leuser National Park were observed to

move from the swamps into the hills to exploit local
increases
in
food abundance."
Social behavior
Orangutans are remarkable among the great apes

in
that they
groups.
appear
cribed as living
to lack distinct social units or
Sumatran orangutans are des-
Instead,
in
loose'
communities that consist
one or more clusters
of
genetically related
of
females and the adult male with which they
all
" Researchers have noted that

movements of community members are subtly
prefer to mate."'
the
coordinated, and that they

Singleton/SOCP
Ian
Fruit
makes up
the
those of females. High-status or dominant adult
larger part of an
males, however, appear
orangutan's diet
small
through most of
nance, within which they attempt to monopolize
the year. Here, a
access
rehabilitated orangutan
subadult lunflanged) males must search for
In
Jambi province,
Sumatra,
is
enjoying a
home range
to receptive
males over
to
maintain a relatively
during their period of domi-
females." " The other adult and
a larger area, usually avoiding the
The overall extent
home range
to
monitoring
influenced by food availability
size
orangutans that do not
as
fruit
gradients.'' In
many
of the
it
are
"
In
rugged, those
is
themselves
restrict
to
Suaq Balimbing, which

fruit
is
largely
flat,
synchronously;
here the orangutans are less mobile on a daily

basis,
but occupy very large
Females occupy home ranges

these swamps, compared
of
to only
home
ranges.'''
around 8.5 km'

3
km'
in
"
in
Ketambe,
while subadult and adult male ranges are around
188
very close
is
in
weakens with age; by
gradually
the time the apes are fully adult, they are mainly
solitary. Interaction
to
among
adults
glances," although juveniles
close relatives." Individuals

related females at
often limited
is
may
play together
home range
in
clusters of closely
Suaq Balimbing not only share
boundaries, but appear to coordinate
their breeding; the timing
of
births
was
similar
within a cluster, but differed between clusters."

After
of soft-
varies over steep altitudinal
major food trees
it
range where they were born and maintain amiable
not clear"
is
circumscribed area follow the availability
pulped
The mother-infant bond

orangutans, but
together as a
''
After independence, females tend to stay near the
movements
Ketambe, where the terrain
may come
occasions.
relations with local females, which are likely to be
and
utan
some
fe-
which Sumatran orang-
of the post-release
group on
areas
occupied by the dominant males.
papaya supplied by one
staff.
real
becoming independent
males move away and either

or
wander over
large areas,
of their
mother,
settle in a large
range
sometimes even well
beyond the forest region occupied by the breeding

population.
As subadults they may
still
travel
together, but flanged adult males generally avoid
encounters with each other
When
flanged males
meet, violent aggressive displays can ensue, and

potentially fatal fights
may occur" Subadult and
flanged males
commonly occupy overlapping home
ranges, however, and flanged males will tolerate
subadult males provided that they keep their

distance."
Long
knit
prominent means by which
calls are a
orangutans
may
maintain links within their loosely
communities. The calls carry over long dis-
tances and
may enable females and
remain aware
of the location of
juveniles to
the males. Only fully
developed adult males produce the long
which
call,
"starts as a series of quiet bubbling grunts, then
builds up into a full-blown gravelly roar
...
often
accompanied by vigorous branch shaking."^ The

throat
act
pouch
inflated during calls
is fully
and may
as a resonating chamber," and the calling
male's hair stands on end during the display Calls

are
made
quency
is
higher^
^^
three or four times a day. Calling fre-
higher where local orangutan density
males may help
of adult
to focus
and direct the
between
long calls, and that cheek-pad differences
Bornean and Sumatran orangutans

gent long
is
has been suggested that the cheek pads
It
relate to diver-
calls.'"
An orangutan community
under the influence
of
*'
attractive females."^'
is
brought together
abundant food and sexually

Sufficient fruit
is
not always
available to sustain groups of orangutans, however,
even should they wish
different
be gregarious.*
to
do aggregate,
individuals
it
is
in
one
When
three
of
Ian Singleton/SOCP
modes:
thought
consortships,
male
in
which a receptive female and
travel together for
an extended
to
have
The mean
period;*'
individuals feed together but leave the food
was about
source separately; and
others for 68 percent of the time
travel
bands,
in
which
all
individuals
feed
be about
1.5; individuals
and 5i percent
in
1.7,
the latter"
Ketambe
it
the former area
in
" Consortships were
longer duration at Suaq Balimbing, where several
of
may
females have also been seen
the next patch together
while at
have been observed with
together within a patch, leave together, and

visit
Suaq Balimbing
daily party size at
has been seen
to
orangutans spend more
use" (see below).
temporary aggregations, which occur when
While Sumatran
transmission of
facilitated the social
skills including tool
time
in
groups than the
Bornean species, they

generally lead solitary
lives,
with males ranging
over a
much
bigger area
than females.
converge on one
to
male, creating a large mating aggregation that also
As single food sources must be divided among the

group,
aggregations are
participating individuals.
likely
to
be costly
By forming these
to
parties,
however, orangutans are thought to obtain mating

opportunities, protection of females
from harass-
ment by subadult males, and opportunities

socialization of infants.*' At
males with mid-sized infants are

participate
ticipate in
for the
Suaq Balimbing,
least
likely
attracted subadult males."

all
attributed
to
These differences are
the greater and
availability of fruit in the
swamp
more
reliable
forests at
Balimbing. which effectively frees orangutans
Suaq
to
be-
have more sociably by relieving the chief constraint

on proximity, that
of food scarcity
and competition.*
feto
Tool use
groups, and adult males rarely par-
Wild Sumatran orangutans have been observed to
aggregations other than consortships."
use 5A different tools for extracting insects or
in
Nevertheless, the high density of orangutans
in
honey, and 20 for opening or preparing
fruits. ^Tool
Sumatran swamp
is
use has been conservatively estimated
to
forests,
and
their sociability,
occur
in
189
World Atlas
Nest building
Sumatran orangutans sleep
new one
build a
nests and usually
in
every evening. They weave together
branches, twigs, and leaves; this normally takes
sometimes takes up
only a few minutes but
to
20
minutes.' About 5 percent of nights, however, are
spent
old nests that the orangutan finds just
in
before retiring,
in
which case the inner
renewed. The main use
of a night nest
lining
is
is
for that
nights sleep but, sometimes, orangutans also play

or rest
in
it
during the following day. Infants share a
nest with their mother until they are weaned.
Sumatran orangutans also

in
which
sometimes cover
when
it
make
often
nests
during the middle of the day. They
to rest
their nests with leaves
and twigs
and have been observed holding a
rains,
leaf or collection of twigs
over their head to provide
shade from the sun, or as an umbrella against

the
rain.'"

Sumatran orangutan males usually reach puberty
at
the age of
Chapter
some mature as
years, although
some
early as five and
as late as
6.
As described
in
the males of both species of orangutan
9,
undergo a complex maturation process, with some
males becoming
fully
mature, or 'flanged'
(i.e.
with
developed cheek pads), earlier than others,
fully
which may remain 'unflanged'
many
for
years.
Flanged males have larger bodies, are more aggresIan
Although the
65 percent
mother-infant bond
are
weakens as orangutans
insects'"
used
mature, females tend to
Neesia
stay close to the range
in fat
which they were born.
in
of
feeding sessions
extract
to
and
cf.
matayana
tfie
Sticl<s
fruits."
seeds
nutritious
Neesia seeds are
of
ricfi
but protected witfiin a large, very tougfi, five-
angled woody capsule. As
tfie
dehisces (bursts open), exposing a

hairs,
among which
sit
ripens,
fruit
mass
it
of irritant
the rows of seeds. To access
the seeds, an orangutan holds

stick
Singlcton/SOCP
Sumatra."
in its
mouth
a short
from which the bark has been stripped, and
rubs the stick inside the
fat-rich aril Ifleshy
first
date
in
in
the
at
well'" (see
190
Box
and not
Suaq Balimbing and evidence
in
the Singkil and Tripa
11.11.
to
has been witnessed

in
the
embedded
Swamps
as
around every eight years."'"

about one month.
lasts
have perineal swellings around ovulation

cycle, but their
marked proceptive
signals their fertile status; that
is,
at
mid-
display behavior
they actively seek
sexual encounters with favored males.
'^
Flanged males usually mate with females

during a consort
sex with
it
reach
Unlike other great apes, orangutan females do not
solicit
of fallen fruits with the tools still
has been seen
to give birth
Their menstrual cycle
seed coatl develops, which the
forests of Sumatra,
typically
years earlier than do males, and are
to
any dryland forests;
many times
form
swamp
thought
five
tends
in
orangutans remove by hand. Tool-use has only been
seen
Female Sumatran orangutans

maturity
the season, a
the capsule. Later
in
and dominant over the others, and are the
preferred mates of adult females.
dislodge the seeds.
fruit to
This technique can be used as soon as the
crack appears
sive
termites and otfier social
access
to
in
be
relationship.
The consortship
by the female.'^ Males actively
initiated
fertile
females
via
'male presenting',
by posturing and displaying the penis. Consortships
may
last
up
to three
months
in
Sumatra and several
females may converge on a single male

in
contrast to the shorter and
sortships
in
Borneo.'"'
more
at
one time,
exclusive con-
" Flanged males are also
known
to force copulations with
rare
Sumatra.^ Consort copulations have a
in
females, but this
is
much
SUMATRAN ORANGUTAN [PONGO ABELII
higher chance of producing young, as the male
her most
with the female during
Females may mate with more than one male during
may
the menstrual cycle, and
more than one
take
years of age
the wild,'" and recent analyses of
in
that they
may
among bonobos, homosexual behavior

has been reported infrequently among great apes
of
the wild.
occurs
only
same-sex
observed
orangutans, such behavior usually
between
captive
or
at
genital contact or manipulation
Suaq Balimbing,
net
loss
Male-biased
of
abundance
some
least
at
in
50.'"
to a relative
males during
areas,
their
dispersal.""
rehabilitated
Over a V 000 hour study, two instances
individuals.
of
In
way
females as adults
suggesting
beyond
in fact live
sex ratios at birth give
in
Ketambe suggest
data collected over 30 years at
cycle to conceive."
Except
may reach 45
Both species of orangutan
is
period.
fertile
were
involving a pair of
Coping with disturbance
Orangutan densities are negatively affected by

selective
but
logging,^'
subadult males that had formed a party together."
assessment
of behavioral
has
there
been
little
change and ecological
The interaction was associated with kiss-squeak'
adaptation
vocalizations (usually an alarm call! and branch
disturbance. Those individuals that try to remain
shaking, which were interpreted to indicate
their
Another interaction between
petitive social tension.
two adolescent males was observed

this
was associated
behavior" There
is
with
too
particular function
com-
more
little
Ketambe;
at
friendly affiliative
evidence to identify any
homosexual behavior
for
in
orangutans.
while
Unflanged subadult males
females
try to associate with
but,
when females
home range
it
is
being logged.
is
apes move. However,
If
if
an
new
regenerating forest
among
Where
but the most
determined subadult males from mating with

her.
Unflanged males do force copulations upon
available females
when
they can," and will follow
females closely for extended periods, interfering

with their foraging efficiency.'"
an infant
carrying
may
find
it
A female
more
that
is
difficult
to
both refugee and resident individuals.^'

the original population
in
any case, even a subadult male
is
this
is
unlogged forest
Puluh National Park.
not at carrying
After a couple of
probably because the area had
decades, this area had
Comparisons
is
of
nest counts
Ketambe
recovered enough to be
in
logged and
suitable as a release site
indicated that orang-
for rehabilitated
utan populations declined by 40 percent, matching
orangutans. Staff
at
the 40 percent decline
pulped
fruit."
area had
Ian
escape;
Selectively logged and
adjacent to Bukit Tiga
previously been logged or hunted.
all
where
cause crowding, stress, and probably starvation
influx of
male and form a consort relationship with him. The

able to prevent
of the
individuals into an area
live,
capacity,
is
most
orangutan populations are
are ready to conceive, they seek out a dominant
flanged male
away from the
dispersal
possible, then
close to the carrying capacity of the forests
they
in
are susceptible to injury or death
logging operations
will
potentially receptive
Sumatran orangutans following
of
in
the availability of soft-
The apes remaining
become more
in
monitor them as part of
the disturbed
the post-release
more.
program.
folivorous, traveled
Singlelon/SOCP
larger
than an adult female. During enforced consortships,

a
female
will try actively to
and seek the protection

available, although
evade the adherent male
of a
she
flanged male
if
one
will not necessarily
is
mate
with him.
The orangutan gestation period
is
about 2i5
days or just over eight months.' Infants are carried

by their mothers for several years and
to
suckle until they are
five or six
may continue
years
old.'
This
gives the orangutan the slowest breeding rate of any
primate, and one of the slowest of any mammal.'^
The young are carried
may
in
a side-ventral position
and
be played with by the mother, and sometimes
share her food. By
11
months
beginning to find their

individuals
of age,
they are
own food." Immature
become independent
of their
mothers
and may wander away from her home range

seven
to 10
at
years of age.
191
World Atlas
Box 11.1 CULTURE AND SOCIALITY

SUMATRAN ORANGUTANS
Orangutans,
appear
chimpanzees Isee Box
like
have
to
either species - that
IN
simple
cultures:
behavioral
easily attributable to ecological distinctions,
are likely to be the result of social learning.
researchers from
When
compared the animals' behavioral
repertoires,
17
be widespread
another,
behavior variants were found

least
in at
one
differences."'
study
considered
Sumatran and Bornean orangutans. Some
(building
and Ketambe
to
with leaves' (holding leaves near the

giving the kiss-squeak
alarm
Borneo but 'tree-hole
of the
that
display
As summarized
shelter an
distinct
cultural
populations. At
in
likely to
in
in
the
Gunung
the north of Sumatra. They
Sumatran orangutan poputwo
of the three populations
mouth while
larly
in
the table below, there are
between the two
differences
use both feeding tools
(at
tree-holes,
scoops'. They also perform 'nest raspberries', 'twig
one Sumatran population.
them
biting
of
twigs
them
into the lining of a nesti,
in
Sumatran orangutans
front of the
in
prior to inserting
and the 'symmetric
scratch' (slow, exaggerated, scratching
of
have cultural origins, observed
passing the ends
movements
two locations
at
Suaq Badmblng
Ketambe
customary'
absent (without ecological reasons!
Seed-extraction tool use
customary
absent (with ecological reasons!
habitual
Branch scoop
Nest raspberry
customary
Twig biting
customary
absent (without ecological reasons)
Symnnetnc scratch
customary
rare
habitual
customarj'
Kiss-squeak with hand

Leaf padding
absent (with ecological reasons!
habitual
Autoerotic tool
customary
Sun cover
habitual
a 'Customary' indicates a higtier frequency of occurrence thian
move on
in
the
to
tool use' (using a twig to
Behavior
forced to
and
get the seeds out of Neesia sp. fruits) and 'branch
Tree-hole tool use
and rested less and
forest,
riverine forest;
were seen only
call!
these cultural behaviors was diagnostic
Behaviors
and
hill
daysl.
swamp
Suaq Balimbing, orangutans regu-
mouth, or actually
of
piling
variants (seven and four, respectively!.
to
biting' (by
were found only
(a
most apparently cultural behavioral
scoop' (using a leafy branch to get water from a tree
None
primary
lations, representing
probe tree holes for insects or honey] and 'branch
hole!
shorter bouts. They were
ground, because of the
discontinuous canopy, which
is
more
energetically
expensive than arboreal travel for an animal that
192
Leuser National Park
occupied nest below from rami and 'kiss-squeak
in
is
a lowland
is
these research sites are both located
of
island. For instance, 'bunk nests'
complete nest above,
shade on warm, clear
for
Suaq Balimbing
both
behaviors were exclusive
apparently cultural
one or the other
and 'sun covers' (orangutans
host the best-studied
comparative
island.
raspberries'
and branches over both the nest and
themselves
to
cultural behaviors."
This
nest
seen and heard as the orangutan finishes
leaves
Each well studied
orangutan population exhibited a unique suite
islands, but absent
one other population on each
building a nest)
no clear, relevant
despite there being
ecological
lips,
but absent at
site,
In
phhhhhp' noise made by blowing through pursed
and
study sites
at least
These behaviors include
the established orangutan
all
one
more populations on both
from
between wild populations that are not
variations
may
be most interesting are the behaviors found

or
i.3|,"
no behavior was universal
is,
on one island but absent from the other What
is
fiabitual'.
so well adapted to
life in
the trees. However,
unlogged long enough, and
area
is left
still
persist
in
if
If
the
orangutans
surrounding areas, densities can
eventually recover
in
the Sekundur area of the
Although only Sumatran orangutans have been seen to develop tool use
tool-use behaviors
of both
arms,
when
something that resembles
in
calis-
thenics or T'ai Chi|. At Ketambe. orangutans use

'leaf
padding' (holding leaves to protect the hands
while handling spiny
stimulate their
to
'autoerotic tools' (a stick
fruitsi,
own
and sun
genitals],
covers'.
Both orangutan populations display the cultural

behavior 'kiss-squeak with hand' (holding a
cupped hand near
mouth while
their
flat
or
giving a kiss-
Over 30 years after Jane Goodall

regular feeding-tool use
in
first
chimpanzees
Suaq Balimbing orangutans were the
so
far,
Primate populations with regular and
show greater social
differences
to
A study of individual
frequency among Suaq
tolerance.^'
tool-use
Balimbing females supports
this
model, showing
spent with other adult orangutans at
less than 50
distance correlates positively with
detailed
comparison
Balimbing
to test this
and
in
measures
initial
of
spent with other orangutans at distances

than 50
two
showed
little
difference between these
orangutans
spent more time
time
of less
detailed analysis revealed that
sites.^''*' Ivjore
the tool-using
to
Bornean
of
in
at
Suaq Balimbing
close proximity to other indi-
and,
close.^'
This
suggests that tolerance
partners at close proximity
is
of
social
an important factor
in
spreading and maintaining regular feeding-tool use
through social learning
in
orangutan populations.
The overall greater gregariousness

orangutans compared
explain
to
those
of
why Sumatran populations
cultural behavior variations.
have
many more
and
learn
from
Sumatran
of
Borneo can help

exhibit
more
Sumatran orangutans
frequent opportunities to watch
one
another,
behavioral
so
of social interactions
model further Orangutans
showed signs
of
five years,
carried
over half the trees
damage. By 2001
Michelle Merrill
at
the 1970s at a removal rate of 11 large trees
per hectare. After
their populations.
Ketambe has been
Gunung Leuser National Park, logging was

out
orangutans yet studied:
known
are
innovations can spread relatively rapidly through
tool use.'"
conducted
Ketambe
be more social than any population
a wider variety of social partners to approach so
(and
widespread feeding-tool use have been proposed
both Suaq Balimbing and
Gombe,
first
the only! wild population of another ape
Suaq
both species adopt
especially, at less than 2 ml; they also permitted
species found to regularly use twigs as probes while
at
vrild,
reported
at
the
that time
the
viduals [both at a distance of less than 10
squeak).''
feeding."
in
in captivity.
still
tree density, fruit
availability,
and orangutan densities resembled
those of pristine forest elsewhere

not
known whether
had recovered
the total
in
number
the park."
of
to prelogging levels, or
It
is
orangutans
whether the
193
World Atlas
remaining orangutans had spaced thennselves out

to exploit available
resources, but
it
is
clear that
the habitat had recovered sufficiently to support the
same number
of
Even less
orangutans as an unlogged
is
plantations.
to
They do enter
plantations or gardens that can provide food, but
such areas are not used

isolated populations in
exclusively.
Sumatra
There are some
surviving
in
rubber
gardens with a very few forest trees along some

of the
stream valley bottoms. These animals are
remnants
The Ketambe River near
its
confluence with the
Alas River,
in
Gunung
panding
alone
it
that
oil
off
by rapidly ex-
palm plantations. Nevertheless,
seems
(some are
have been cut
still
severely fragmented forest, as their fruit-
in
dominated
ranges
their
main ecological
dispersers. Virtually nothing
about this aspect
Sumatran orangutan ecology,
of
Herman
intact seeds. This topic
and
is
to
survive long
Serge Wich
discussed
in
Chapters 2
10.
Our limited knowledge about their frugivory
ecological role of
populations are probably unable
by
Rijksen," kU percent of which contained
breeding], although they are grad-
Sumatra.
be as seed
known, however,
is
beyond the 96 fecal samples investigated
left
exterminated as pests. Most orangutan
the orangutan diet,
fruit in
role is lil<ely to
and seed dispersal makes
ually being
occupy large
to
Ecological role
Given the importance of
they might survive at low densities
Leuser National Park,
them
requires
diet
ensure sufficient supplies."
to
forest.
known about how Sumatran
orangutans cope with shifting agriculture and

conversion of land
term
to
it
define the
difficult to
Sumatran orangutans
in
relation
other species. Orangutans share their environ-
ment
with several frugivorous species of similar or
larger size, such as elephants [Etephas maximus],
rhinoceroses [Dicerorhinus sumatrensis], sunbears

[Helarctos malayanus], and,
of
Lake Toba
species
in
may
in
forest patches south
These
only, tapirs {Tapirus indicus].
be less important than the orangutan
seed dispersal,' but practically nothing
about their role or the degree
is
known
of dietary overlap
with orangutans. Elephants and rhinoceroses are
able to transport
much
larger seeds internally than
orangutans can and tapirs are considered good

seed dispersers in neotropical forests,"' " as are
sunbears
in
Borneo.'^ Of the other primates, the
siamang [Symphalangus syndactytus], which

30 percent the size of an orangutan,
capable of dispersing
many
of the
is
is
15-
also probably
same
seeds.
interactions with other animals
The food resources
of a forest
sympatric animal species

ecological
when
niche to
to
are limited, forcing
feeding from the large strangler figs that
attract
great diversity of animals,
orangutans usually share most

fruit
own
develop their
reduce competition. Except
species
with
primate species.
In
around
of their
five
Sumatran
preferred
medium-sized
any given lowland
forest,
are a subset of the following: pig-tailed
these
macaques
{Macaca nemestrina]; long-tailed macaques

fasciculans]; five leaf
spp.,
monkey species
IM.
[Presbytls
Trachypithecus spp.l; and three gibbons
(two small species of Hylobates and the larger
siamang].^" These primates all

fruits,
canopy, but differ

All but the
194
consume
and leaves; they are active

in
siamang
in
figs,
other
the forest
food selection and range size.

will feed peacefully with
each
SUMATRAN ORANGUTAN IPONGO
Table 11.1 Feeding strateg les of sympatrjc apes
Time
Symphatangus
10-13
territorial
territorial
0.4
0.2
2-10+
05.00-06.00
06.00-07.00
07.00-08.00
very fast
fast
very slow
range Ikm'l
of rising
Speed
of travel
Distance/day Iml
250-1 500
consumed
Quantity of food
Ketambe area
6-8
Ikgl
Territoriality
Home
the
Hylobates
Species characteristics
Weight
in
small
Ecological strategy
Pongo
30-70
overlapping
home ranges
400-800
500-900
medium
very large
least selective feeder; stores
selective feeder;
aggressive
prefers small sweet figs"
competitor
Fruit (percent of dietl
56
52
60
Leaves (percent
34
40
34
of dieti
excess energy as
Adapted from Rijksen, H.D. 119781
/Pongo pygmaeus
other
the
in
same
tree, in
Ketambe, female and
abelii
Field Study on
primate species and usually feed peacefully
siamang, but apparently only by certain individuals.
roosts periodically, causing a
Ketambe orangutans usually show no
to
hesitation
in
that allow
them
to
consume more mature
leaves than other primates do. Although they are
same hard-husked
able to process the
fruit
orangutans, they avoid the sweeter pulpy

favored
as
in
trees
their
influx of bats
in
trees
these trees during the day, which
can deter orangutans from entering them, even

in
the case of a fruiting
Arctlctis binturong]
ing trees
Bearcats (binturong,
fig.
show
a tendency to avoid fruit-
where orangutans are feeding."
Humans appear
fruit
by orangutans, gibbons, siamang, and
mass
in
may shift
an area. They establish feeding territories
and may remain
monkeys possess complex, fermentative
Leaf
stomachs
tissue
Sumatran Orangutans
with them. Fruit bats [Pteropus spp.]
where siamang are feeding."
fat
Lesson 18271 H Veenman and Sons. Wageningen.
young orangutans have been seen being attacked by
entering trees
to
be the only frequent
predator of adult orangutans, but the latters near-
macaques. " Orangutans are also distinguished
exclusive arboreality
from the other primates by their greater tolerance
dation by tigers [Panthera tigris sumatrae].^"
of unripe or acidic fruits."
body
The two macaque species found

from
differ
all
in
- in their ranging behavior. Long-tailed

live in
rivers,
Sumatra
other primates - and from each other
large groups
small
in
macaques
home ranges
where they remain unless
close to
fruit
is
very
that
of
may
reflect the risk of pre-
an old male orangutan was found
appeared
to
The Sumatran
have been freshly
tiger itself
Endangered on the Red
List of
The
1975
killed by a tiger"
designated
is
in
Critically
Threatened Species,
with only a few hundred individuals remaining

200i.
hunting dogs ICuon alpinus]
huge home ranges and travel long distances on the
immature orangutans." There are no leopards
in
forests
away from
large rivers.'"
"^
in
Clouded leopard [Neofelis nebulosa] and
abundant elsewhere; pig-tailed macaques have

ground
ABELIl]
the species Panthera pardus
may
in
occasionally take
of
Sumatra or Borneo.
The orangutans and gibbons occupy the most

similar ecological niches, but
ensure competition
still
have features that
minimized (Table
is
11.11.
The
smaller gibbons [Hylobates spp.l are more selective

feeders, rise earlier
in
the day, and remain within
small territories (usually 0.5 km^ or
less),
which they
defend against conspecifics. Orangutans consume

large quantities of fruit at
excess energy as
fat.
one time and store the
Siamang show intermediate
characteristics."
Orangutans show
HABITAT REQUIREMENTS
Sumatran orangutans are most abundant
freshwater and peat
they
may
response
to
non-
the
swamps
(Table 11.2],
10 individuals per square kilometer.'' They
intermediate densities
and
hill
where
occasionally reach densities as high as
in
live at
lowland dipterocarp forest
dipterocarp forest, and can be found at very
low density
little
in
forests of flood plains, alluvial bottomlands, and
forests."
in
some submontane and montane

in some secondary
Orangutans also occur
195
World Atlas
Table 11.2 Sumatran orangutan densities
Habitat type
Flood plain and peat
Ihill
lorest
and dipterocarp
forest!
Submontane and montane

a
Mean orangutans per km^,
In
foUov^êd in
in
6,9 14.5-7,01
3,213,0-5,51
1,211,1-1,31
1,1
0,7 10,4-1,21
the
in
3-5 max,
7+)
11,0-2,21
1,0
3 (max. 61'
parentheses by information on range, wfiere available,
the Upper Alas valley
forests.
Selectively
orangutans
logged
areas
relative to undisturbed
parable forest composition;
tfie
support
areas
of
few
com-
reduction depends
tfie
to
be determined by two main factors - the avail-
intensity of the logging that has tal<en place."
The current
of
distribution of
preferred fruits and
Hunting by
humans may be
complete absence
of
presence.
responsible for the
orangutans
in
some
regions of
Sumatra, and their continued decline parallels that

increasing
human
of
low
10
occupation and clearance of

Ian
Singleton/SOCP
human
C.P.. eta/.
population density.'" Rivers
cm deep
wide and 60
120011"
These apes survive best
their forest habitat.
in
areas
more than
also restrict orangutan
movement.
orangutans seems
human
and van Schaik,
Rijksen, H.D., luleijaard, E, I1999P'
on
of
Density^
Leuser Ecosystem"
Density
From
ability
multiple locations
of locations
swamp
and bottom land
Secondary and
Upland
different habitats
Density"
No
Alluvial
in
Orangutans prefer soft-pulped

availability of
such food
orangutan density;"
is
fruit
and the
an excellent indicator
of
explains low densities at
it
high altitudes' and
in
also their variable
abundance
recently logged forest," and

specific habitat
in
more important
types. ^' Certain fruit species are
than others. Lianas
make up
orangutan food plants
(fruit
17
percent of
and leaves]
in
all
Ketambe
South of the provinces of
and they also provide an important means
North Sumatra and
arboreal transport.'" Strangling figs produce large
Aceh, the only free-living
fruit
orangutans
food supply for the apes,
are
in
in
Sumatra
the environs of
of
crops at short intervals, providing a regular
Sumatran swamp
where they
occur, in
some
forests, species of Tetramerista,
Bukit Tiga Puluh
Sandoricum, and Neesia
National Park, Jambi,
where over 50 ex-
A tropical forest can be conceived as an

"immense patchwork of plant communities of
fulfill
this function.^'
captive orangutans have
different compositions. "''
so far been reintroduced
usefulness
by the Sumatran
richest patches are
peat forests, between rivers, and
Programme, and are
area surrounding these habitats. These areas have
now
breeding.
to
Each patch varies
in
its
an orangutan seeking food. The

in
swamps,
floodplains, and
in
the 10-15
km
high diversity of productive food trees;" 30-50
percent of trees typically provide
orangutans and 10 percent
fruit
suitable for
of the trees fruit
each
month. However, low-quality patches for orangutans
occur within these otherwise good
still
habitats.
percent
It
of
has been estimated that perhaps 50
an expanse of
swamp
forest constitutes
suitable habitat for orangutans."

in
rivers
comparison, the drier areas away from

are
dominated
by wind-dispersed
dip-
terocarps, with few regularly fruiting trees. Only
around 35 percent
196
of
these areas are thought
to
SUMATRAN ORANGUTAN IPONGO ABELIl]
form suitable orangutan habitat." During dipterocarp mast fruiting events that occur every few years,
orangutans move
advantage
these dry regions
into
masses
of the
to
take
seeds that are
of fat-rich
produced. Further discussion on dipterocarps and
mast
fruiting
Above
given
is
Chapter
in
10.
000 m, Sumatran orangutan densities
decline markedly, although occasional pockets of
higher density occur

Individual
upland valley habitats.
in
wandering' males are seen as high, or
higher than
500 m, but breeding populations are
These
not supported."
altitudinal
limits
mainly
reflect the distribution of favored food types: that
many
the upper limit of
whose
trees
The healthy orang-
type preferred by orangutans.'
utan populations found

of the
in
the two high plateaus
Alas Valley are thought
to
The
depend on the
an unusually high
less acidic soils that support
abundance
is.
fruits are of a
required
support
viable
habitat
The questions
swered about
also
make
ranges
for
and
that
it
difficult to
typically
hill
It
needed
is
assumed
for
needed
in
it
precisely^'
Home
km^ or larger
The
largest
incorporate areas of
swamp
for females.'""
higher
that
viable
its
food-rich areas.
in
250-500 individuals are
population," which would
require between 50 and 600
depending on
be an-
Range overlap varies considerably
habitats and
is
to
and ranging systems
5-25
of
home ranges
forest."
remain
still
estimate
males and 1-10 km'
among
to
their social
are
occupied
of figs."
area
Sumatran orangutan population varies with

quality.
km'
of
occupied forest,
Larger areas are
suitability."
practice, to allow for local patchiness
in
orangutan distribution.
It
is
possible to estimate
how many orangutans would be supported
in
100 km' of forest of quality similar to that of Suaq
POPULATION STATUS, TRENDS, AND THREATS
Logging and mining,
Human
such as here,
Paleolithic
human migrants may
have settled along
in
and
adjacent to Tanjung
Balimbing, assuming that ranging patterns are also
Sumatra's east coast and larger rivers as long as
Puting National Park,
the range sizes and degree of overlap for
80 000 years ago. Based on archeological evidence
Indonesia, have had a
the Padang Highlands of western
devastating effect on
similar"
If
each age-sex class
yield a density of 7.25/km',
then
100 km' would support a total of 725 orangutans. Of

these, 229
would be adult females and 100 would
be subadult males, but only 33 would be
from caves
in
Sumatra, these people consumed orangutans
Seven hunter-gatherer peoples are known
fully adult
expected to be dominant
Sakai, Akit, Lubu,
at
any one time, and thus
blocks
20 500 km'
enough
of
in
primary
forest
area. Only 9 000
altitude to sustain
totaling
km' was
and Ulu." These peoples
mainly on the eastern side
By 2002, Sumatran orangutans occupied only

13
around
at a
low
permanent orangutan
to
Asia.
Sumatra; Abung, Kubu, Mamaq,
have existed
in
orangutan habitat
throughout Southeast
relatively large quantities.'"
males. Of these 33, fewer than seven would be
contribute significantly to the gene pool.
in
of
lived
the Bukit Barisan
mountain range, occupying the banks and dry

ridges of the extensive floodplains and peat
of
swamps
eastern Sumatra south of Lake Toba." The fact
that few,
if
any, orangutans survive
in
this region
populations." Only four blocks supported over 500
today suggests that they exerted heavy hunting
individuals (see below).
pressure on the species.
197
World Atlas
fed
upon an orangutan
liver
and then sacrificed."
Various other items were also adorned with orangutan skin or hair; even as late as
in
the Alas Valley
971 a Batak youth

,
was seen wearing an orangutan-
skin cap.'"
Recent history
During the 20th century, orangutans began to be
viewed as having an economic value, and a considerable International trade occurred
and
960s.
in
the 1930s
The Dutch professional animal
van Goens, captured
collector,
218 adult Sumatran
at least
to export to circuses
and zoos abroad.
The Gayo people were renowned
for their ability to
orangutans
trap live orangutans.
Numerous
infants
were also
sold into the national and international pet trade,
many
with
being exported to Taiwan.
Orangutans continue
to
be killed by local and
foreign people, despite legal protection.

still
an Smglelon/SOCP
A Sumatran orangutan
before confiscation
forests of the
in
December 2003, found

in
and
a cage at the back of a
restaurant
in
Desa
Riau province. He
Petal,
was
his
embedded
abdomen.
this
lil<ely to
is
in
in
historic times,
persist there.
to
much
have inhabited
people
Local
the
be the reason that great ape
Muslims who do not eat orangutans
centuries.
from
absent
Leuser Ecosystem
populations were able
wildlife!
later found to have air

rifle pellets
were
Hunter-gatherers
the
of
area for
timber
extracted
some
subsistence needs and caused
Devout
most other
(or
for
habitat dis-
Islam to
Achenesel has halted their hunting

for food.
of all faiths
elite
organized hunting safaris
as late as the 1990s."
and Meijaard
In
appear
to
have been the most effective hunters.
Apes were among
their favorite prey
"somewhat sweet, but
nice
hunted with dogs and spears
in
Antiahs toxicaria
source
system
in
to their
they were
the lowland parts of
and poison-tipped
The darts were often poisoned with the sap
of the
fruit
due
taste";'
their range, or with blowpipes

darts.
Indonesia
Sumatra
of the
of the
the higher
tree,
ironically a favorite
orangutan. The traditional belief
hunters deterred them from hunting

hills,
where
spirits
were thought
to
orangutans
people
in
army
northern Sumatra
the recent past Rijksen
towards
orangutans - that they have "an economic, medicinal, nutritious
central
of
killed by
aptly described the feeling
warrants persecution.""
of
is
they raid fruit crops'" and to obtain
if
primarily agriculturalists and
The hunter-gatherers
still
infants as pets. There are also reports that the
impact on orangutan populations.'''
the fates of
or nuisance value - any of which
is in
all
a unique historical position,
its
and
wild species will be greatly
influenced by the outcome of complex social and
economic processes over the next few years. The

country
means
is
reinventing
clear.
itself,
but as
what
by no
is
There are powerful forces exerting
various kinds of influence, including pressures for
and participatory democracy opposed by
local
others
who seek
democracy.
a return to centralized 'guided'
Political, religious, military, capitalist,
and bureaucratic
elites
compete
for influence
opportunity at the local, regional, and national
and
level.
Indonesian development
reside.^' Certain clans or families also traditionally
The meteoric pace
avoided eating orangutan meat, but did not criticize
during the 1970s to 1990s can never be repeated, as
their neighbors for hunting orangutans.^'
The Batak people

utan meat will
make them
into modern
shamans were
strong, and this belief
has prevailed
times.
wands
often
of
orangutan
hair""
it
believe that eating orang-
The
staffs
and
decorated with
Other reports claim that
to
draw
the spirit into the staffs, a kidnapped child had to be
198
However, the apes are
turbance through shifting cultivation, but were

had a very limited
The meat
some in Sumatra, but the spread of

many traditional people (Gayo, Alas, and
eaten by
was
of
fuelled by the opening of virgin lands
clearing of virgin forests that have
used up, by the sale
now
of
and the
now been
all
but
petroleum resources that are
depleted, and by the borrowing of
wealth that has largely
fled the
immense
country leaving a
legacy of debt. The chief issue for orangutans and
most other species
of
the
Sumatran
forests
is
"
SUMATRAN ORANGUTAN IPONGO ABELIl]
momentum
whether or not the habits and
model
lished by this
sustainability are
Indonesian people.
Pongo
Ecosystem finding
Union.
way
the Leuser
of
(with the support of the
nongovernmental
Indonesian
and
organizations,
to
practical terms, the fate of
In
depends on the peoples
afae//)
European
addressed by the
last
at
their friends
government]
in
secure the ecological architecture
own
of their
environment and, thereby, a future for themselves

as well as their neighbors and cousins
(see
Box
The Leuser Ecosystem
estab-
will destroy the
fundamental issues
frontier, before the
remaining
of
development
of
is
in
Leuser Eco-
system, which includes Gunung Leuser National

Park, has been the focus of European Union support
Leuser Development Programme Isee Box
via the
However,
11.21.
into
this
ecosystem
least four areas
at
is
now fragmented
Leuser,
forest (West
of
Trumon-Singkil, East Leuser, and Tripal, with the

prospect
new
of
further fragmentation due to a proposed
road scheme."
Recent models propose that populations
to
The current position

It
the best-
of
of
over 500 Sumatran orangutans are large enough
11.21.
number
what
1900, and 2 percent of
ago when the
it
Sumatran
of
what
less than 7 percent of
is
be demographically stable
1993,
estimated that the total
orangutans
one
range. The
heart of their present
the forest
in
is
protected sites for Sumatran orangutans, and the
was
was
it
10 000 years
It
was thought
that
the long term."
in
six
existed; by 2002, four remained, only
was
one
outside the Leuser Ecosystem.
of
which
addition,
In
were thought
eight smaller populations that
In
such populations
to exist
to the
south of Lake Toba had disappeared by 2002;
and Sumatra became isolated from the Asian
two
these have been
mainland." " The decline of the species accel-
and the others may
erated towards the end of the 20th century, with
1993" Isee Table
massive exploitation
occurring
the
in
age ended, sea
last ice
Sumatran
of
1970s,
continuing to date.
absence
the
human
of
would be
intervention, the recovery of this species
extremely slow (around 0.006 percent per year] due

to their very
slow reproductive
lost before
Sumatran orangutans are unable
to survive
long term
in
in reality
]^.A].
severely fragmented forests. Habitat
fragmentation, and degradation are the major
loss,
threats to orangutan survival."

of the
in
Sumatran orangutans."
immediate
population
extinction
risl< of
future.'
fragmented
is
is
It
into a
in
the
clear that the
number
Table 11.3 Sumatran orangutan population decline since 8000
in
extinguished.
unknown.
are
turn,
their
being
Year
Estinnated
number
Decline from
of
Sumatran orangutans
of
split
previous estimate
small
-
380 000
up or
1900
85 000
78 percent
1997
12 500
85 percent
7 334
41 percent
The exact population sizes are
Rijl<sen
and Meijaard estimated that

2001-2002
Sumatran orangutans survived
in
1993, and
"
predicted that by 2020 only 7 500 individuals would
remain. This
BC
the wild
8000 BC
that,
12 500
In
as
classified
2000, indicating that the species
faces an extremely high
units
The fragmentation
Leuser Ecosystem probably has the greatest
Endangered by lUCN-The World Conser-
Critically
vation Union
the
over the 10 year period
lost
have been
single impact on
rate.*'
Sumatran orangutans were
in
of
forest habitats
and 1990s, and
1980s,
In
levels rose,
From
Ri]k5en H D
Meijaard.
E 119991." and S ngleton,
etaf, eds 12004]
was based on the fact that 45 percent

was formally protected." This pro-
of their habitat
jection
now considered
is
The 1993 figure
of
to
have been optimistic.
12 500
orangutans was an
overestimate,
because
populations
southern Sumatra which are
thought
to
in
it
have been
to
now
decline since 1997.
000 individuals have been estimated

lost
to
each year between 1997 and 2000
from the Leuser Ecosystem alone." By 2002, only
around 7 300 individuals are believed

remained" Isee Table
Table 11.4 Estimated populat ion structure of Sumatran orangutans
unconfirmed
have already died out at the time.
Populations have continued

At least
included
to
have
Surviving population size

< 100 individuals
1993"
2001-2002"
10
100-500 individuals
500-1 000 individuals
>
000 individuals
23
Total
From Rijksen
H.D., Meijaard, E. 119991.^
13
and S ngleton.
et
a(.,
eds 120041
.3|.
199
World Atlas
Box 11.2 HISTORY OF THE LEUSER ECOSYSTEM

During the 1980s, the work of
Rijksen. Mike
Gunung Leaser National Park
the proposed
7 927
Herman
Carelvan Schaik, and others revealed that
Griffiths,
km' was inadequate
sentative
sample
made
of
northern
Sumatra. This should not have been a surprise, as

the boundaries, set
in
1936 and 1976. bear
little
One boundary section was simply

a
was thought
it
line
the time
of
designation.
boundary was a semicircle
of
of
Proper demarcation
is
condition for
national park
weaker
in
of
such as elephants,
990s revealed
and hornbills. Similar
fruit bats,
studies revealed that species such as the orang-
The
forests to maintain viable populations.
full
initial
itself.
gazettement
boundaries
it
of a
has a
failed to
take into consideration the natural ecology of the
bounded area
naturally
same
the
size
was named
samples
Combining
of
as
Haiti
of
27 000 km' - roughly
or Rwanda.
This zone
Leuser Ecosystem', and includes
'the
most Sumatran ecosystem
types.
The Leuser Ecosystem stretches from the
radius,
boundaries on the
Indonesia, prior to which
legal status.
the
Another
km
30
centered on the summit of Gunung Leuser
ground
with the geomorphology of the region revealed a
and two important
was an inadequate knowledge

at
the early
in
wide-ranging Sumatran species
of
not to be significant
tops 50
water catchments were bisected by the boundary,
geography
Studies carried out
movements
the
the habitat needs of these charismatic species
that a unique volcanic plateau
or there
be resolved before the national
to
km
map between two mountain
apart. Either
would need
park could be legally formalized.
utan and tiger needed extensive tracts of lowland
relation to the terrain.
drawn on
of
the placing of physical
boundary markers practically impossible - an issue

that
conserve a repre-
to
and fauna
of the flora
at
mountainous character
region. Furthermore, the
the planned park
sandy beaches bordering the Indian Ocean,
right
across the breadth of Sumatra almost to the
man-
swamps
grove
bordering the Malacca Straits.
It
includes two great mountain ranges reaching over
3 000
rift
in altitude.
These are separated by a great
which two large
valley through
rivers flow - the
Tripa to the northwest and the Alas to the south.
The Alas
River,
one
of
two major rivers flowing
through the Leuser Ecosystem.
The
rivers flow into the Indian
Ocean, after passing
through extensive freshwater peat

are
home
Earth.
is
It
to the
swamps
that
densest orangutan populations on
the last place where viable lor potentially
populations of the Sumatran varieties of
viablel
elephant, orangutan, tiger, and rhinoceros exist,
and the only place where
all
these species are
found together
was
It
only
identifying the
pleted
in
after
the
scientific
work
in
Leuser Ecosystem had been com-
990. that
it
was
realized that
most
of this
area had already been promoted for conservation

by the traditional leaders of the peoples
part of
Sumatra as
local leaders
ernment
early
1960s,
Sumatra was almost completely
covered by tropical forest, but this cover
was
greatly
reduced and fragmented by logging, infrastructure
upsurge
in
that
had lobbied the Dutch colonial gov-
have the forests of Leuser conserved
in
deforestation due to the collapse of
control by central
government
in
Indonesia, follow-
development, resettlement (transmigration), and
decades of poor governance and environmental

neglect." " Kalimantan (Indonesian Borneo! was
plantation development on a massive scale during
also affected, and the impacts on Indonesian wild-
There was a 61 percent
the 1970s and 1980s.^
'"
decline
Sumatra between 1985 and
in
forest area in
1997." From 1998 onwards there
200
to
in
early as 1928. For six years, the
was
sudden
ing
life
are immeasurable.
Given that 80 percent

habitat
is
of
current orangutan
covered by timber concessions, or
is
vul-
SUMATRAN ORANGUTAN {PONGO ABELIl]
Eventually an area approximating the
perpetuity.
Gunung Leuser National Park was granted,
ex-
cluding nnost of the valuable lowland forests and

coastal plains.
Once the Leuser Ecosystem was

an area
tremendous significance
of
conservation management, the great chal-
holistic
was
lenge
as
identified
that required
to translate this vision into reality.
effort required to
do so has proved
was one important
but there
easier than
it
to
made
factor that
might otherwise have been.
early 1990s, the
The
be enormous,
European Union was keen
In
it
the
to help
conserve tropical rainforests and the Indonesian
government was keen
was
there
to
solid financial
do the same, as long as

support from the inter-
national community. This meeting of policies culmi-
nated
a series of
in
commitments
by the European
Commission and the government

provide about
tions
US$38
place to conserve the Leuser Ecosystem.
in
This commitment, which began
be completed toward the end
form
2004.
Project
for
to
These
which was
first of
new management system
for the area
was needed because
in
develop a
The second important achievement was
as a whole.
includes
to build
support for the conservation
Ecosystem among an array

central and
plantations and locally claimed lands, as well as
business
natural forests under various kinds of planned use.
others.
This
meant
that decision
Station in the Leuser
Ecosystem.
important
to
area
the
Aceh,
Ketambe Research
Programme.
several
yielded
activities
achievements, the
Serge Wich
took the
It
Lowland Rainforests
followed by the Leuser Development
This
was due
1992,
in
of
an Integrated Conservation and Develop-
of
ment
of Indonesia to
million to put the right condi-
making was fragmented
The
governments,
local
community,
third
was
local
to gain
and conservation management capacity was also
the
weak. The government
presidential, ministerial,
assigned
a
management
private
foundation,
Indonesia
of
of the
the
therefore
Leuser Ecosystem
to
Leuser International
Foundation, which had established
itself
with the
sion
in
all
national level.
required 3 000
provide
support through the
Leuser
and provincial
with
management, and
The consolidation
km
of
boundaries
fires
continue at high rates. The
and droughts that ravaged Kalimantan over the
past two decades have been less of a threat to

the orangutans on Sumatra, so
far.
There, approxi-
mately 5 percent of the 1997-1998
occurred within orangutan
of the
its
inclu-
to
concrete markers every 2
be demarcated
km
task
continued overleaf
nerable to illegal logging and habitat conversion,

is likely to
have
of this legal status
International Foundation.
habitat loss
the
at
levels
plans from the local to the
spatial
express aim of conserving the Leuser Ecosystem.
to
its
and
legal recognition for
been issued supporting the legal status
Leuser Ecosystem,
the
universities,
communities,
Leuser Ecosystem. Various decrees
The Leuser Development Programme then continued
Leuser
of the
stakeholders from
of
fire
hotspots
habitat." Large-scale
forest fires to the south of the
orangutan range
area,
in
in
June
control
were
central Sumatra, were reported
2004. Urgent consultations on
underway between the
smog
(Malaysian, Indonesian,
and
Singapore governments.'
The tsunami
of
December 2004
the tragic destruction of
that led to
many towns and
around Aceh's coastline had
little
villages
direct impact on
orangutan habitat. The indirect impacts remain
to
201
World Atlas
requiring massive investment of labor and funds.
Those charged with the work had

through the surf
the
of
cement bags and
night temperatures
markers on
the
installed
They had
isolated coastlines.
where
Ocean, carrying
Indian
steel rods as well as supplies to
them as they
sustain
row ashore
to
climb high peaks
to
hover around 0C.
Another, and hopefully the

lenge
network
roads called Ladia Galaska
promoted by the province
some
through
Leuser: a
of
of
being
is
Aceh. This would pass
of
the biologically richest forests
Leuser and would open up access
in
logging,
for
In
plantations, and settlements (legal or otherwise).
addition to the physical challenges, the demarcation
The Leuser International Foundation has developed
was done during
national and international alliances to oppose this
crossfire.
armed
a time of
and there was a
was
to
conflict in Aceh,
the
plan and a decision on the future of Ladia Galaska
was
that few local
had been expected shortly after the presidential
lengthy dialog
elections
real risk of being caught
further challenge
people understood conservation.

therefore needed at
in
the settlements near
all
the planned boundaries before the boundary
markers could be erected with
The
bottom-up
signing
before provincial leaders and finally
the central government
was
in
Jakarta. The whole task
eventually completed, however, and the Leuser
Ecosystem was formally constituted
ments
at all levels
as an
legal docu-
in
conservation area.
official
The foundations had thereby been
for
laid
the
zonation and protection of this priceless area.
Not
all
virgin forest.
areas had been given out for
logging or even plantations, and
some were
little
if
anything to improve
alternative transport
arrangements would do
harm and
more
contribute
The tsunami
December 2004 had few
on the Leuser Ecosystem, with only
direct effects
the Tripa
of
swamp
temporary
area
influx
known
to
have experienced
seawater" Following the
of
devastation caused to Aceh's coastal communities,
some
m^
people have suggested that the 4-8 million

logs
of
being
be imported as a form
have called for the timber
to
sourced from sustainably
managed
forests
in
temperate countries."
to convert
lowland forests to
accelerated or delayed as a result
settle
people from Java
effort.
to
work on
Leuser
estates.
to agricultural land.
Hence an ongoing
Foundation
International
Development Programme has been
Leuser
resolve
to
these conflicting land uses. By June 2004, more

than half
of the
12 original togging concessions
had been closed down, and the licenses

problematic plantations revoked.
has stopped and there
anywhere
in
be seen, but
it
is
Swamp
of several
drainage
At stake ultimately
is
the long-term future of
the inner core of the Leuser Ecosystem - about

21
000 km', mostly
will
of forest. This area,
safeguard the supply
ment services
to
of
feared that the need for firewood

is likely to
lead to further
likely survival of
preserved,
and the many other denizens
of Leuser's forests.
use by researchers since then;

the Alas Valley within the
it
Sumatran orangutan
in
Renun^' and Rijksen
Ketambe.'Only Ketambe has been
1993
in
swamp
is
Griffiths
located
in
forest at
was opened
Suaq Balimbing
in
the
south of the Gunung Leuser National Park. These
behavior and ecology were carried out

at
It
the orangutan
Park. Another long-term study site
MacKinnon
if
water and other catch-
4 million people downstream.
would also mean the
in
1970s, by
not
the rebuilding
of
Mike
detailed studies of
is
no cattle ranching
loss of forest.
first
is
It
project will be
Leuser
and construction timber
The
whether the Ladia Galaska
clear
job for the
and
be
of international aid, being
Plans also existed
to
should
required for rebuilding
swamps
ranches and
less
development.
to local
drained to convert
cattle
Inter-
sourced from the Leuser Ecosystem area.' Others
Leuser Ecosystem comprised
the
Some
road project would do
Ikabupaten] leaders
affair with district
off first,
dif-
demarcation. This was a
physical
September 2004. The Leuser
in
national Foundation will continue to argue that the
the overall welfare of local communities, and that
local consent.
process was even more
ratification
than
ficult
202
of
great chal-
last,
conservation
the
facing
IS
in
the
at
continuous
two
sites,
in
environments
lowland and
with
relatively
exceptional
orangutans, have yielded almost
about
the
all
Sumatran orangutan
fruit-rich
densities
that
in
is
the
of
known
wild,
SUMATRAN ORANGUTAN iPONCO ABELIl]
some
although
work has also been
additional
done by Priatna
Soraya and Sikundur, and by
at
'
Fox at Agusan.'
The Sumatran Orangutan Conservation Pro-
gramme
toring
conducts most
survey and moni-
of the
work concerning the status and
of wild
Sumatran orangutans
at
established a release program
National Park
in
distribution
present.
has
It
Bukit Tiga Puluh
Jambi province. The Research,
in
Monitoring, and Information Division of the Leuser
Management
manages research
Unit
activities
within the Leuser Ecosystem (see Box 11.21.

it
Initially
ran four research stations:
Ketambe, which
is
operational (lowland
still
habituated
with studies focused on
forest),
Thomas
and
orangutans
monkeys,
leaf
Presbytis thomasi;
Soraya (lowland
protected within a log-
forest],
down as
ging concession but closed
a result of
An orangutan
the conflict;
Suaq Balimbing (lowland peat swamp

closed
down as
a result of the conflict;
Bengkung (based
in
and
moved
release site
a transmigration site in
in
to its
Jambi
province.
down
the middle of lowland rain forest), closed
as a result
being
forest),
the conflict.
of
Two monitoring posts have also been established,

considerably extending the potential for survey work
and diverse Leuser Ecosystem."
within the vast
Whether any
remains open
maneuvers
of
this
work
to question, for
will
be enough
these are our
final
preserve the Sumatran orangutan
to
in
Awareness
the wild. After centuries of population contraction
and decades
of deforestation
animals have nowhere else
to
and logging, these
orangutan's plight
is
of the
being
taken into schools by the
go but the already
fragmented Leuser Ecosystem and one other forest
Sumatran Orangutan
patch nearby. The best estimate of the current
Conservation Programme.
population
is
about 7 300, and
it
is still
believed to
be declining despite considerable conservation

investment
in
and around the Leuser Ecosystem.
Nevertheless, the concentration of opportunities for
Sumatran orangutan protection may
itself
make
the
strongly synergistic.
people,
learn
how and why
forests, then
and enforcement efforts
orangutan
close proximity can be
breathing space can be

efforts,
during which
governments, and businesses can
local
task easier, as consen/ation, research, education,

in
If
maintained through these
it
is
to
preserve enough of these
just possible that the
will survive within
Sumatran
them.
FURTHER READING
Corlett, R.T. (1998) Frugivory
and seed dispersal by vertebrates
In
the Oriental llndomalayani region. Biological
Review73: itU-Ua.
Djojosudharmo,
Sumatran
S.,
van Schalk,
C.P. 11992)
forest. Tropical Biodiversity
Why
'I
(1):
are orangutans so rare
in
the highlands: altitudinal changes
in
11-22
203
World Atlas
Knop,
Ward,
E.,
Wich, S.A. 120041 A comparison of orang-utan density
P.I.,
Sumatra. Biological Conservation 120
MacKinnon,
J.
The behaviour and ecology
119741
UtamI, S.S., Goossens,
of wild
forest
on
orangutans IPongo pygmaeus]. Animal Behaviour 22: 3-74.
Sumatran orang-utans. Behavioral Ecology ^2
in
Ancrenaz, M., Borgen,
C.P.,
and unlogged
Bruford, M.W., de Ruiter, J.R., van Hooff, J.A.R.A.M. 120021 Male bimaturism and
B.,
reproductive success
van Schaik,
a logged
in
187-192.
121:
G.,
Galdikas,
643-652.
151:
CD., Singleton,
B., Knott,
I.,
Suzuki,
UtamI, S.S., Merrill, M.
A.,
(20031 Orangutan cultures and the evolution of material culture. Sc/ence 299: 102-105.
van Schaik,
Fox, E.A., SItompul, A.F. 11996) Manufacture and use of tools
C.P.,
implications for
van Schaik,
human
Monk,
C.P.,
Robertson, J.M.Y. 120011 Dramatic decline
K.A.,
Ecosystem, northern Sumatra. Oryx35

WIch,
S.A., Singleton, L,
Utami-Atmoko,
Sumatran orangutan Pongo
abelii:
111:
VKild
Sumatran orangutans:
S.S.,
orang-utan numbers
in
in
the Leuser
14-25.
Geurts, M.L., Rljksen, H.D., van Schaik, C.P. 120031 The status of the
an update. Oryx 37
(11:
49-54
WIch, S.A., Utami-Atmoko, S.S., Sella, T.M., Rijksen, H.D., Schiirmann, C, van Hooff,
Sumatran orangutans [Pongo
history of wild
in
evolution. Naturwissenschaften 83: 186-188.
abelii].
Journal of
Human
J.A.,
van Schaik, C.P (20041
Evolution il
161:
Life
385-398.
MAP DATA SOURCES

Map
11.1
RIswan (2004) Orangutan
DadI, R.A.,
Leuser Management Unit as part
distribution polygons: developed at the
Leuser Development Programme, funded by the European Commission and the government
Leuser Management
field data:
Priatna,
J.,
Unit,
Sumatra, Indonesia. Based on technical
van Schaik, C, Idrusman, Singleton,
D.,
Rijksen, H.. Riswan, Robertson,
Bloxam, C, Kuswandono. Long.
Y.,
1.,
Wich,
I.
Main sources of
Additional information from DadI,
R., Griffiths, M.,
criteria set by Singleton,
Bogor Expedition
Universities of Bristol and
McPherson,
B.,
S.
and members
J.I,
of the
of Indonesia.
of
the Leuser
to
Sumatra (Burton,
Management
Unit's
Antipoaching Unit.
Singleton,
R.,
1.,
Wich,
Byers,
0.,
S.,
Husson,
S.,
Stephens,
S.,
Utami Atmoko,
S.,
Leighton, M., Rosen, N., Traylor-Holzer,
eds (20041 Orangutan Population and Habitat
Viability
K.,
Lacy,
Assessment: Final Report lUCN/SSC

ACKNOWLEDGMENTS
Many thanks
to Raffaella
Commitante
Morrogh-Bernard lUniversity
Rondang Siregar
The orangutan
of
lUniversity of
lUniversity of Cambridge!,
Simon Husson
(University of Palangkarayal, Helen
Palangkarayal, Ian Singleton ISumatran Orangutan Conservation Programmel, and
Cambridge!
distribution polygons
for their valuable
comments on
were produced by Rahmadi
A.
Dadi and Riswan
of
the Leuser
Management
Unit
as part of the Leuser Development Programme, funded by the European Commission and the government of
Indonesia. They
were based on technical
criteria
designed by Ian Singleton,
In
turn based on
many years
of field
research on the Sumatran orangutan (especially on densities by habitat type and altitudinal limitations], together
with the most recent Information from the Leuser
Management
was
who
were: Caret van Schaik, Idrusman,
digitized
Ian Singleton,
most
of the forest-cover data.
The main sources
was
Yarrow Robertson, members
Leuser Management
Kim McConkey, UNEP World Conservation Monitoring Centre

Box
11.1 Michelle Merrill,
Box
1 1
Griffiths,
Bogor Expedition
James McPhersonI, and members
Unit.
AUTHORS
Mike
greatly assisted by Nick Jewell,
provided by Rahmadi A. DadI, Mike Griffiths, Dolly Priatna,
of the
Catherine Bloxam, Kuswandono, Barney Long,
.2
of field data
and Serge Wich.
Additional information
20/1
Unit on destroyed or degraded habitat Ifrom
extensive ground surveys, aerial photomosaics, and satellite Images!. This team
Emergent Systems
Leuser Management Unit
of
to
Herman
Rijksen, Riswan,
Sumatra (James Burton,
the Antipoaching Unit of the
Gibbons: the small apes
Chapter 12
Gibbons:
the small apes
David
The
some
great apes of Southeast Asia share their
range. Apart from clarifying distribution
family's
and abundance from lesser-known areas,
apes: the gibbons (family Hylobatidael.
lysis is
cases, the gibbons are even
attention. This brief overview
intended
less public
to raise
the gibbon family, and offers the
of
profile
is
In
more threatened
much
opportunity to contrast their unique behavior and

of the
ecology with that
gibbons
in
of the family
field in
the
shown
to
Thailand
in
now
divided into four genera.
Thomas Geissmann (w\Aw.gibbons
These are mainly
del
lar or
white-handed
gibbon [Hylobates
larl
v/ith infant.
study of
the 1930s,' most species
Hylobatidae were
950s,
are
DNA ana-
the key to resolving disputes. The gibbons
other apes.
Following Carpenter's pioneering

lar
Chivers
habitat with smaller but no less interesting
than the great apes, but receive
the
J.
970s, and
be monogamous,
first
studied
the
in
980s. They have been
territorial, frugivorous,
and suspensory, with elaborate duets by the adult

pair
The complexities
investigated
gibbons
in
of
each species have been
recent years, and the
both seed
dispersal and
roles
of
forest
re-
generation have been demonstrated. Gibbons
live
in
on the mainland
of
Southeast Asia, and on the
islands of Java, Sumatra, Borneo (Kalimantan), and
associated islands. These
on the Sunda shelf,
all sit
which emerged from the sea as a consequence

volcanic activity about 12 million years ago Imyal.
owes
its
admixture
of
It
uniquely rich fauna and flora to an

of
immigrants. These
came
first
from the
Indian subcontinent (the Siva-I^'lalayan fauna]
and
then later from China (the Sino-Malayan fauna).'
The gibbon lineage diverged from that

apes about 15 mya somewhere
in
of the
other
forested, tropical,
or subtropical Asia.
GIBBON TAXONOMY
There are
validity of
still
burning
issues concerning the
species to be resolved with regard
gibbon taxonomy, especially
in
to
the northeast of the
205
World Atlas
large
Reus tree
In
the
riparian forest along
the river KInabatangan,
Borneo. FIcus are

critical
resources for
wildlife since they
produce
fruits several
times a year.
allopatric Iwith disjunct ranges) except wtiere the
siamang range overlaps
that of the lar
and
2.
H.
pileatus (the
gibbons.
3.
H.
moioch
now
HI
Symphalangus, the siamang,

of
S.
syndactylus,
4.
Sumatra and the Malay Peninsula (two
Nomascus, four species
of
"
" " from
5.
H.
of Java,
white-handed gibbon),
agilis
(the
in
agile
in
in
Thailand
the Malay
north Sumatra:
or black-handed
gibbon), with
Laos:
range stretches from the Malay Pen-
2.
3.
N.
concolor (four subspecies) and N.
leucogenys
N.
in
center
the
in
insula (between two lar subspecies) to
Sumatra south
(two
of
in
the rest of
Lake Toba, and one
the southwest of Borneo (West and
subspecies):
in
N. gabrieilae in the south:
Central Kalimantan, bounded by the
Kapuas and
Bunopithecus,'^ the hoolock gibbon, B. hoolock
Myanmar
into the
6.
subspecies radiating around the rest
(extending across northern Thailand
southwest corner
Hytobates, comprising
of China):
five
to
six
Barito Rivers); and
H. mueileri (MiJllers gibbon), with three
and
species,
In
the Sunda shelf:
the last two groups
view of the extensive hybridization between

in
the center of Borneo,"
H. klossii [the Kloss gibbon), confined to
may
the Mentawai Islands
as a fourth subspecies
of the latter, but
argues that the agile
is
of
Sumatra:
off
of
Borneo."
ranging from Thailand through the islands of
1.
one subspecies whose
the east of Sumatra, one
the north:
(two subspecies), of Assam, Bangladesh, and
206
H. lar (the lar or
Peninsula, and one
crested gibbons,
nasutus (two subspecies)
(4)
Javan gibbon),
southern China, Viet Nam, Cambodia, and
1.
13)
(the
land Yunnan, China), one
each with several subspecies,"
of
confined to the west:
with two or three subspecies
subspecies):
(2)
gibbon),
pileated
southeast Thailand and west Cambodia;
agile
the west coast
it
be necessary to sink H. muelleh into H. agilis
more
Geissmann
similar to the lar
gibbon." He also defined the four genera, based

on molecular data suggesting a
as tar bacl<
split
The
calls
and pelage color and markings are
among
monochromatic
the gibbons.' Species are either

the west
(to
and
black,
southeast grey], polychromatic
more open semideciduous
the
to
the center], or
(in
asexually or sexually dichromatic

the
Habitat
Gibbon habitats span the two main forest forma-
as 8 mya.'
distinctive
ECOLOGY AND BEHAVIOR
lin
the north,
in
habitat); this reveals
tions of this part of the Oriental biogeographical

region:" the
semideciduous monsoon
evergreen rain forests

the islands of the
forests of the
of the
Sunda
Sunda
Malay Peninsula and
shelf."
shelf
The evergreen
habitat. Significant
numbers
pelage coloration have a genetic basis; these
viduals occur
more seasonal
features, with profound behavioral significance for
land Asia, concentrated
reproduction, should therefore be taken seriously
forest,
The other key parameter
species and sexes

call'
is
in
for
song, with the female 'great
in
the
and survive
maritime influence
Thailand,
in
in,
rain
comprise the main gibbon
an intriguing geographical pattern. Both calls and
classifying gibbons.
rain forests of
'mainland Asia' north of the isthmus of Kra; and the
of
species and indiforests of main-
in
pockets of evergreen
the
moister areas under
for
example,
Indochina,
Myanmar. and Bangladesh.
being the diagnostic feature. Family groups of
gibbons tend to sing daily to advertise their territory
and the strength
of their pair
bond. Male and female
gibbons have distinctive parts;'^
among
a true duet
is
it
the majority of species, which
more common
primates, although
in
most unusual
is
in birds.
GIBBON EVOLUTION
Chivers has proposed a
relating to frequent
model
changes
of
sea level during the
As
at high
Gibbons and
orangutans eat
ice
exposed as one land mass; as the

the shelf
formed
and altitudes, the Sunda shelf was
latter part of the Pleistocene.'
latitudes
gibbon evolution,
of
was
flooded, a
number
ice
many
melted and
of islands
were
evolved
and
separation,
in
then
(left)
migrated once land bridges were restored. The key

point in this
model
is that,
occurred
Sunda
the
within
Ibelow).
to
Serge
V\/ich
gibbon speciation
shelf,
with
shelf
back
sequential spread
and Ficus sp.
after the initial spread of
three ancestral lineages lor gibbon general into

different parts of the
same
such as
Blumeodendron sp.
exposed. The wholly or partially isolated gibbon
populations
of the
fruits,
left
the
subsequent
mainland. This
competes with the idea that gibbon species spread

out from the Asian
model,
Chivers
the
mainland. According
hoolock
gibbon
to
the
was
the
enter the Asian mainland, followed by the
first to
pileated
and
lar
gibbons; the Kloss jMentawail,
Bornean, and Javan gibbons originated on the
edges
in
of the shelf;
and the
agile
and
lar originated
the center of the shelf. During the periods of
lowest sea level, the center of the shelf dried out; at

that time
the key rain-forest relicts, into which
gibbons and other forest animals retreated and out

of
which they spread when sea
level rose,
were
in
eastern Indochina and southern China, northeast
Borneo, west Java, northern Sumatra, and southern
Myanmar, as well as the Mentawai
Islands.
207
World Atlas
and the Kloss, pileated, and hoolock spend around

70 percent. About 25 percent of the
figs (nearly
40 percent
in
intake
fruit
is
siamangl. Young leaves are
important for most gibbons, especially the siamang,

but
not
Kloss gibbon
the
for
where the
Islands,
Mentawai
the
in
poor and the leaves
soils are
better defended chemically Animal matter, mostly
provides an
invertebrate,
important source of
animal protein (about 10 percent
The underlying preference

sources
of ripe (sugary)
and pulpy
The gibbons, then, are

like
of feeding time).
smaller
the
for
is
(fleshy] fruit.
fruit-pulp specialists,"
compete more than
the chimpanzee, but they
most primates with large birds such as pigeons and

and sugary
hornbills for small, colorful,
food focus
the
seems
location
to learn
resources and
food
The small area
hence the gibbon's
many
territoriality
monogamy The focus
handed gibbon
on small
[Hylobates
large multimale, multifemale groups of
(art singing.
Thomas Geissmann [www gibbons
del
[Macaca
Trees of the family Dipterocarpaceae are
most gibbon
typical of
43 percent
some
and averaging
persed by several bird species, they are less
Leguminosae
important.
abundance from 13
to 14 percent.
trees of the
in
Between 24 and 50
families
tree
documented from gibbon
have
been
Ifigsl
commonest
tree
Euphorbiaceae
are
the
families used as food sources
in
gibbon habitats,
In
dispersing seeds, animals encourage
same
Gibbons
differ
by foraging
of
gibbon biomass.
45 percent
known gibbon
of all
prefer lowland forests,

of fruiting trees
species, the
in
at
and density
in
has a greater
the diet and occurs
more
in
activity
through the heat
Gibbons spend between 57 and 72 percent
lar,
and flowers). The siamang, which
208
and
agile
of the
day
they retire early for the night, usually several hours

before sunset. This frees food sources shared with
other primates,
langurs [Presbytis and
Trachy-
pithecus spp.] or macaques [Macaca spp.], that are
monopolized during the morning by gibbons.
Population density
The density
(in
of their
feeding time eating the reproductive parts of plants
at
the day, and a
the cooler lower levels of the canopy;
of
monogamous
family groups of gib-
varies from 1.5 (for two species
an exception
in
bons, typically comprising around four individuals,
higher-altitude forests.
Feeding and activity patterns
Ifruit
not
about 10 kg, weighs
as the others],
tolerance for leaves

frequently
diversity
greatest; however, the largest
siamang (which,
much
twice as
is
where
Gibbons
foods.^*
in
long midday siesta.'" After active bouts of feeding,
gibbons sustain
good predictor
from other primates
with feeding peaks early and late
Rubiaceae, Clusiaceae, and Anacardiaceae. Figs are

a particularly
asynchronously,
fruit
availability
having a markedly bimodal pattern of daily activity
followed by Leguminosae, Myrtaceae, Annonaceae,
These families present 161 species, comprising
species to
which prolongs food
habitats (averaging 371,
with about 400 trees per hectare.' t*/loraceae
and
plant species, gibbons are key seed
dispersers; for others, especially those also dis-
from
16 percent,' whereas the tree family

varies inversely
macaques
and the large-bodied orangutan.
to
habitats, ranging
of forest composition,^*
For
competition with the
fruiting trees avoids
spp.)
and
monogamous family groups
Adult male lar or white-
by
to
that can be effectively
protected does not provide enough food for

individuals,
This
fruit.
have required gibbons
suitable
of
protect them.
to
is
larger,
is
44 percent; the moloch, Muller's,
gibbons spend around 60 percent;
in
Malaya]
to 6.5
Thailand) groups per square kilometer Biomass
density, as indicated by gibbon
kilometer,
is
biomass per square
more useful measure
density than the
number
per square kilometer, as
of population
of individuals or
it
relates
more
groups
closely to
food availability, presumably at times of scarcity.'
The combined biomass

in
gibbons
lar
for
Kalimantan and 10^ kg/km'
for
Mullen's gibbon
lar
siamang and
of
Malaya was 126 kg/km^ with 3U kg/km^
gibbons
in
Thailand. Therefore, gibbons are at
in
numerous
least as
more seasonal
the
in
forests
among
the flexible supports of the small trees of
the understory. Indeed, they escape from the heat
midday sun by foraging
of the
fruit
new
or
story.''
'^
leaves
in
Day range
trees, in the high forest
siamang
Pileated and Muller's gibbons, and
from 0.8
km
to 0.9
have been seen

as
much
to travel
and
1.2
as
as
little
km. Siamang
1.5
0.
the
gibbons for the main canopy and emergent
away from edge
habitats.'"
travel
on average, while other
daily
gibbons travel between
for
in
example, shows the prefer-
Malay Peninsula,
of
small trees with
Comparison with other primates
farther north.'
ence
in
the relative cool of the under-
km/day and
as 2.86 km/day. The day range of the
Social organization
Group
size
averages 3.8 - equivalent
to
and two young - but ranges from two

there
may sometimes
an adult pair
to seven,
i.e.
be three or four young.' Only
hoolock gibbon varies from 0.28 to 3.A0 km; other
the concolor gibbon has been recorded as living
gibbons show comparable variation, from about
polygynous groups, with two or three adult females
0.40 to 2.50 km.
These figures
food distribution.
In
reflect variation in
the Malay Peninsula,
ranged less and ate more leaves when
when
increasing day ranges
scarce,
abundant and energy
monsoon
forests,
levels
siamang
in
size of 7.2 in
Bawanglin Nature Reserve,
were
Hainan,'" although this requires confirmation. Infants
were
are aged up to two or three years until wholly cap-
fruits
fruits
excess.'
in
and young, and an average group

Yunnan'^ and 5.3
in
In
the
able of independent travel; juveniles up to
five
or six
however, where leaves are not
such a viable alternative

increased day ranges
for the
may
smaller gibbons,
Gibbons compete with
wider search for
large birds such as
reflect a
hornbills for their
sufficient fruit.'
preferred sugary
Home range and territory

Home range varies between
Thailand and
in
to 0.45+
km'
km'
0.56
0.1 7
for
for lar
home ranges
km'
for
0.16
in
for lar
gibbons
moloch gibbons
hoolock gibbons
gibbons
km'
in
fruit.
in
Java,
Bangladesh and
Malaya, where siamang
Where
same area
are also large (0.3-0.4 km').
there are two species of gibbon
in
the
lone always being the largest species, the siamang],

it
that the home ranges of both are larger

when alone because of competition for
is likely
than
particular fruiting trees. Agile gibbons have smaller
home ranges
that are evenly used over any five day
siamangs and
period;
home ranges
with
lar
gibbons each have larger
more
both
limited
and more
variable patterns of ranging over five day periods,
same fruiting tree

home range that
although centered on the
The proportion
trees.'"
defended as territory
of the
for the exclusive
use
or
is
of the
resident group varies: siamang, 62 percent; Kloss,
64
percent;
hoolock,
86
percent;
Muller's,
88
percent; moloch, 94 percent. Other species defend
75-77 percent
Vertical
of their
home
ranges.
use of forest canopy
With their suspensory behavior, gibbons exploit

the high forest
forest
canopy more than most other
primates,
but they are
equally at
rain
home
209
World Atlas
independently Overt
expressions are limited
facial
open-mouth threats
to
aggressive/submissive
in
interactions.
Only
the siamang does the male carry the
in
infant during its
weaned from
second year
the female
of
after
life,
(although
has been
it
may
it
suckle at night as
it
infant learns first
about those animals on
is
most dependent
sleeps with her),
still
way
the
whom
it
for its survival: the female, the
male, and then the subadult with

while the adults groom.
whom
juvenile, with
in this
whom
plays
it
Interacts least with the
It
one might expect
to
it
play
most. The adult female usually leads the group
around the home range, hence the need

carrying
growing
the
infant
subadult lags behind at the rear
from
that the adult male,
influencing the direction

Thomas Geis&rnann Iwww gibbons. del
move between
front as they
clear,
however,
travel.'
is
The smaller
forage on a broad
to
the main food trees.
Grooming involves either adults and subadults
gibbon [Nomascus
during rest periods, or adults and young as they
gabhellae).
Right:
of
is
It
central position,
Its
gibbons separate more often
yellovz-cheeked crested
to stop
earliest
The juvenile follows the female, the
opportunity.
Above: Juvenile male
the
at
tends to sleep with
settle for the night [the juvenile
Female and male
the male, the infant with the female). Play
the
is
slamang [Symphalangus
other main social
syndactylus\; an adult
of the active
pair during a duet, with
pileated,
partially Inflated throat
spend much time playing alone - swinging, jumping,
sacs.
recorded
activity,
day
some
In
in
up
to
4 percent
studies (siamang,
and hoolock). While the
infant
lar,
and juvenile
manipulating tree parts - they also swing from,

grapple with, and bite
at,
adults or subadults, and
sometimes other juveniles.

Singing and social interactions between groups
The duet
tain
in all
mate and
availability
gibbon species
territory,
believed to main-
is
specifically to
and attract a mate,
to
advertise
develop the pair
Thomas Geissmann Iwww gibbons, del
bond (and cement other bonds within the group),

years; and subadutts, physically adult-like, to eight
years or so,
when
and
to
seem
they leave the natal group.
defend the mate and the
to
and males
Social interactions wittiin groups
Social interactions
virithin
groups are
relatively
infrequent, presumably because the family group
so cohesive and so familiar
vs/ith
its
is
follow their parents.
The only sounds heard, apart
from the resonating group

of
calls
foliage, are
to
exclude strange males to defend their
These songs are reinforced by bound-
ary patrols while food
chases
and the movement
squeals from an im-
to
and
fro
The complex
makes
It
difficult to
Interaction
in
Borneo
'
and on
ently to the
songs
it
is
encouraged
to
move
multiple factors
in
playback experiments on Mijller's and agile gibbons
too close to a parent (usually the male), or the bleats

distress as
of
simple terms. These have been best clarified by
pair has
in
being sought and during
explain the observed behaviors
The resident
an infant
is
across the boundary.
mature animal, usually a subadult, who has come
of
210
branches and
Females
forest space.
daily routines.
Overt signals are rare, as the young watch and
territory.
exclude other females to defend their mate,
lar
gibbons
been shown
of
to
in
Thailand."
respond
neighbors than
to
differ-
those of
strangers; the former they expect, and the latter
cause much greater
The female reacts
agitation.
strongly to a strange female. Groups will duet
ponse
res-
in
lone female calling, but will approach
to a
silently a lone
male
gibbons respond
that
Bornean
calling.
is
to calls of agile gibbons,
from Sumatra, but not usually
to
those
Mullers
territoriality - that
of the key
monogamy and
sociology -
features of gibbon
are thought to confer benefits as
as imposing costs.
vi/ell
monogamous and
being
In
gamous, the male could be said

potential reproductive success;
to
it
mono-
be reducing his
is
thought
percent of the active day Such
to
be able
weaken
to
Given the stability of gibbon family groups over long
be
to
it
is
rare to observe dispersal of maturing
young and the formation

that has
of
new groups. The
emerged from observations
Mentawai'"
is
of
territory with or
Daughters tend
to
wander
from the parental
less far
sons and are more
leads to this sacrifice, as a result of the energetic
parental help.
costs of patrolling and defending a territory with a
to take over the natal territory
believe that gibbons are
field
workers
monogamous because
they
are adapted to surviving on small fruiting trees.
It
has been argued by van Schaik and Dunbar that
monogamous
gibbons are
and that they could
live in
to
prevent infanticide,
polygynous groups; this
analysis was, however, based on incomplete and
rare alternative
parents disappear;
may
occur,
but
if
this
and/or reproductively
is
likely to receive
for a
young adult
when one
or both
one parent survives, mating

incest
is
usually transient
ineffective.
Thomas Geissmann (www.gibbons
del
Adult male agile gibbon
[Hylobates
agilis);
dark variant with
light
cheek patches.
disparate data."
Some
pattern
Malaya' and
without parental help, and then obtaining a mate.
territory than
and predictable food supply. Most
in
young adults, recently excluded
from the natal group, acquiring a
the available niche and distribution of food that
rich
the basic
territorial pattern.
Group formation
periods,
gibbons.
There has been extensive discussion
occupied
circumstances seem
agile
even those
of
mon and
exceptions to a strictly
monogamous
pattern have been observed, but typically
in
extreme
circumstances. Palombit studied siamang and lar

gibbon
Park
in
much
Ketambe in the Gunung Leuser National

Sumatra." " He observed the pair bond to be
at
stronger
siamang; the pair was more
in
cohesive and equal amounts
of
grooming were
performed by the male and female. Several observed
mate switching, and
cases of mate desertion,

extrapair copulations
dence
of disease
in
or isolated
among
the groups of hoolock
Bangladesh, because they were restricted

in
forest
fragments
of varying size.'
tendency toward promiscuity or even polygyny

relate
to
inci-
and death. Ahsan has reported
similar observations
gibbons
were caused by a high
shortage
of
males
in
Any
may
such forest
fragments.
Reichard and
Sommer echo the argument that
females are defending their mate and males

defending the territory's resources, suggesting that
extrapair copulations 112 percent of those seeni
help to confuse paternity and forestall infanticide:
hence, kin relations extended
into
neighboring
groups.^"'" They studied isolated lar populations
in
Thailand, near the hybrid zone inhabited by both H.
brand
of lar
H. pileatus. Here, the
home ranges
of pairs
gibbons overlapped by around 6^ percent.
Encounters between groups were therefore com-
211
World Atlas
managed
ships of
need
forest areas
to
be planned
carefully on the basis of systematic research,
needs
of
which
of
the strategy
still
use
to
is
much
be conducted. The third part
to
maximum
to
efficiency the
land already cleared of forest or so degraded that

role
its
as forest cannot be redeemed.
Logging, primates, and people
represents
logging
Selective
but
work
will only
timber extraction
is
very
this
approach
in
if
Tekam area
the Sungai
Peninsula.""'" Even
extracted
wliite-cheeked crested
the total stand
gibbon [Nomascus
IS
damaged during
leucogenys teucogenys).
is
the larger and
(i.e.
if
(i.e.
68 percent of the plant biomassl

access, felling, and extraction.
more frugivorous species
fully within
disturbance).
Malay
only 10 trees per hectare are
most vulnerable, but
recover
of the
A percent of the trees), 45 percent of
Adult female northern
the
the long term,
in
and carefully controlled. Johns studied
it
light
compromise
the
between human and animal needs
It
that are
their populations should
20-30 years
(if
there
is
no further
gibbons and
For example,
langurs
adapt their foraging strategies by eating more leaves

as
fruit availability
declines
Gibbons maintain their

affects
their
newly logged
in
territories,
forest.
but the stress
Langurs may emigrate
breeding.
temporarily from the disturbed area, and there
(because of travel
adds
to the
Selective
across gaps), which
difficulties
breeding
is
among immature monkeys
Increased mortality
loss.
logging
enhances the
diversity
of
microhabitats characteristic of the mosaic of successional stages of the forest;
plants
of
immature
is
it
that
forest
the colonizing
provide
more
and less chemically defended foods. Bird
Juvenile male northern
nutritious
white-cheeked crested
communities maintain much the same trophic
gibbon {Nomascus
structure, but species composition
teucogenys leucogenys).
markedly: dietary generalists survive better than

Thomas Geissmann Iwww.gibbons.del
insect and fruit specialists,
GIBBON CONSERVATION
Forest clearance
is
also undermines
human
many
other animals, and
wellbeing. Once the forest
below about
cover of a tropical country dips
and logged forest can maintain viable populations

of
the
large wide-ranging
persistence of primary forest

cial to the survival of certain
Thus, the
hornbills.
in
an area
may be
animal species, and
cruit
is
50 percent, climatic changes and water and soil
the relationships between these two types of forest
problems seem
that
countries
seem
to escalate catastrophically'As
able to afford to keep
few
more than
10 percent of their forests totally protected, at least
another AO percent
of forest
for sustained yields of a
Managed
forests,
area has to be
wide variety
of
managed
products."
forests provide a buffer zone for protected
which provide replenishment
of plants
and
animals. The shapes, sizes, and spatial relation-
212
whose food supply may
be highly disrupted temporarily Mosaics of primary
the greatest threat to the
survival of primates and
may be changed
need
to
be investigated urgently Additional
information on the effects of selective logging
is
available for the Malay Peninsula from surveys of
primary and variously disturbed

East Kalimantan."
In
forest'^
and from
contrast to the tolerance of
gibbons and langurs, orangutans and proboscis
monkeys [Nasalis
selective logging.
larvatus] are seriously affected by
been practiced
Shifting cultivation tias
Where population
rivers.
in
densities are low, there
are long fallow periods between incidents of

cultivation
and
fire
any given place, allowing forest
at
Under these
recovery.
nnuch
Asia for centuries, especially along
of Souttieast
may
conditions, a landscape
be maintained Indefinitely under a mosaic of forest
patches
of different stature
and age, with
little
im-
pact on biodiversity or ecological functions relative

to
When human
natural conditions.
become
density rises, fallow periods
population
so short that
becomes unsustainable.
the system
can
Forest
populations' are being
realized,
with the
Kloss,
moloch, and concolor gibbons being the most
endangered. As the clear-felling
of
forest areas
declines, however, their prospects are boosted
if
adequate selectively logged forest (with low extracas gibbons have shown them-
tion rate) persists,
selves to be very adaptable to such disturbance."'

Little
progress has been
made
niques for translocation - the
groups from
doomed
presumably because
involved,
and the lack
of
to
In
movement
physical
of social
habitat -
a protected
the
""
developing tech-
difficulties
of suitable destination habitat
progress see Cheynes work].'
quickly disappear from the landscape under these
(but
conditions.
remains a possible solution where populations
Human needs mean

managed
that
much
forest
must be
for a great variety of forest products, not
just timber.^''
" What
required
Is
the Improved
is
become
paration,
lowland
ecosystems (especially the
all
with
ones),
management
the
efficient,
of large buffer zones,
richest,
sustainable
and the more
real
critically
endangered, but adequate pre-
care (with veterinary superA/ision), and
Captive breeding worldwide provides valuable

publicity (about the plight of rain forest animals!
and education, with fund-raising opportunities

conservation activities.
It
productive use of land already cleared of forest.'
gene
Such
pects of reintroduction
strategy should ensure that viable popu-
lations of all gibbon taxa survive

will not
be easy
in
perpetuity, but
it
to put into effect.
more
developed
tvlather"^^
analyzing gibbon food
from
trees
all
The pros-
the wild are gloomy,
habitat
is
available.
It
is
of
much
cost effective and successful to translocate
Serge Wich
approach
valuable
the
to
however, given the costs involved and the lack

If
for
also helps to conserve the
pool, with meticulous studbooks.
available habitat.
Translocation, captive breeding, reintroduction
It
monitoring are essential.
protection of watersheds and national parks rep-
resenting
for
of
previous
Degraded habitat
in
Borneo.
studies for comparison with the density of gibbons

in
each area.' He shows that there
pondence
between
abundance
of
more
analysis enables one to assess
of
human
and
the
trees. This
a gibbon
below (because
or above
disturbance!,
fig
whether
at carrying capacity, or
is
a direct corres-
these preferred gibbon foods. Group
size increases In localities with
population
is
biomass
gibbon
(because of
immigration from nearby disturbed areas). The
proposed
suitability of
sites for reintroduction or
translocation can be assessed, and stocking density
determined; where there

reduction
In
is
selective logging, the
carrying capacity can be determined.
Our improved taxonomic and socioecological

understanding
of this diverse
of their habitats in tropical
group
of
apes and
moist forests Improves
the chances for their effective conservation. Clearer

recognition
of
species and subspecies, and im-
proved quantification
structure, feeding,
Is
or
of
the use of resources (social
and ranging]
In
relation to
what
available are essential to effective protection and/
management.
The predictions
of a drastic reduction in
gibbon
213
World Atlas
doomed
social groups from
understocked protected
must be
to protect
wildlife within
fragments
forest
to
prime
Ttie
forest.
Threatened gibbons
any
Indochina
effort
(it
the Bukit Baka National Park
in
crested gibbon species
in
beam of hope. Facilities

accommodate confiscated
and
all
contains about eight of the
20 most endangered primates
it.
Kalaweit
key to gibbon conservation, indeed to
is
primate conservation
natural habitat and to conserve
the world). The four
in
[Nomascus
southern
spp.) in northern
China
Central Kalimantan offers a
Indochina
are being developed to
threatened, but the most endangered are the Hainan
ready!
them
reintroduce
to
Another possible area

developed nearer
to
and (when
pair formation,
gibbons, to support
which
in
to
do
[Nomascus
Palangka Raya, the provincial
flourish.
is
many
in
countries
demon-
are
lives
the world
critical,
however,
decision
makers
ments
is
over
of
of
rapidly to
have
avert
this
if
An
irreparably
not
this
to
be changed
be
genus.
of the
or silvery gibbon
[Hylobates
the west of Java,
in
[H. klossii]
on the Mentawai
numbers
cause
is
for
Bangladesh and
in
eastern India are not large. The pileated gibbon (H.

status
is
restricted
unknown
in
Cambodia. Otherwise, the more widely
H.
in
its
is
siamang [Symphalangus syndactylus],
and Bornean gibbons
lar, agile,
critical role to
Thailand and
pileatus]
distributed
bio-
muellen] are present
forests remain, even
this process.
in
[Nomascus
and Cambodia
Myanmar, and perhaps
in
serious concern;
damaged
interdisciplinary
environmental approach could have a

play
unknown
network concerned
international
disseminating
with
is
and
hoolock gibbon [Bunopithecus hootock]
the
national commercial
impending catastrophes. Values
planet
Nam
Viet
Islands off the west coast of Sumatra. The status of
user countries' as well as the
to
be the most numerous
to
and the Kloss gibbon
be changed and resource flows significantly
to
altered
all
effort local
gibbon
moioch], which survives only
Iwho now mostly see
concerns. Policy and activities need
in
Nam
southern Viet
in
loss are the Javan
These are the govern-
international and
more
The other gibbons most endangered by habitat
be done) and, more importantly, the
to
governments
heads
seems
Most
influence the
to
yellow-cheeked
the
gabriellae]
next
Ithe
essential.
is
need
the
of today.
of tropical countries
what has
while
most immediately affected by
destruction of forests) and the young
generation)
ensure that they
to
are, the
Laos {Nomascus leucogenys] are also struggling,
people
strate. In the long term, education of local
(whose
made
The rarer they
southern white-cheeked gibbons
essential at various levels, as
successful programs
Nam) gibbons
people are often willing to make. The northern and
capital of Central Kalimantan.'^
Education
(northeast Viet
Vit
seriously
nasutus], with fewer than 20 individuals
each. Efforts are being
being
this is
and Ca
(China)
protected forest.
to
are
in
[H. lar, H.agilis,
and
good numbers where
in
selectively logged ones.
FURTHER READING
Brockelman,
H..
W.Y.,
Srikosamatara,
S.
(198A1 Maintenance and evolution of social structure
gibbons.
in
Chivers, D.J., Brockelman, W.Y., Creel, N., eds. The Lesser Apes: Evolutionary
In:
Preuschoft.
and Behaviourai
Biology.
Edinburgh University Press, Edinburgh, pp. 298-323.

Carpenter, C.R. (1940)
field
study
in
Slam
Comparative Psychology Monographs 16
of
the behaviour and social relations of the gibbon [Hylobates
(51:
lar\.
1-212.
Chivers, D.J., ed. (19801 Malayan Forest Primates: Ten Years' Study
in
Tropical Rain Forest.
Chivers, D.J. (20011 The swinging singing apes: fighting for food and family
in
Plenum
far-east forests.
Press,
New York.
Brookfield Zoo, The
In:
Apes: Challenges for the 21st Century. Conference proceedings. Chicago Zoological Society, Brookfield,
Illinois.
http://www.brookfield2oo.org/contentO.asp7pagelDs773. pp. 1-28,
Gelssmann,
T.
(1995) Gibbon systematics and species identification. International
Leighton, D.R. (19871 Gibbons: territoriality and

R,W., Struhsaker.
T.T.,
eds,
monogamy.
In:
Primate Societies. University
Smuts,
of
B.B.,
Zoo News
1*2:
65-77.
Cheney, D.L, Seytarth, R.M., Wrangham,
Chicago Press, Chicago, pp. 135-U5.
ACKNOWLEDGMENTS
Many thanks to
David Woodruff [University of California, San Diego) for his valuable
comments on
AUTHOR
David
2U
J,
Chivers, Wildlife Research Group,
Department
of
Anatomy, University
of
Cambridge
World Atlas
Conserving the
great apes
TOSHISADA NiSHIDA
conservation of nature can be divided into
The
long-term
short-term, middle-range, and

perspectives.
A short-term
aims
project
to
things.
Anthropocentrism
pushing the great apes
we do
not succeed
the
is
force
driving
to extinction. Therefore,
if
educating people to abandon
in
secure immediate protection by actually preventing
our current anthropocentrism there can be no hope
harmful acts through effective law enforcement. On
of saving the great apes.
simple
level,
short-term
project
important than a long-term one, since
more
is
animals
if
Of course,
killing
we humans cannot
are extinct today, there will be no need for a long-
them and should
refrain
term project tomorrow. However, even
seek pleasure or
to satisfy
if
you suc-
ceed
in
saving animals this decade, but then lose
them
in
the next, the current venture
Our mission
pointless.
in
is
completely
the short term should be
subsist without
animals and plants. However, we can respect
from
killing
them
solely to
our excessive appetites.
Only 50 years ago, for example, the prohibition on

of
human
culture, except for rare special occasions
such as
wasting food was a universal tenet
to
save the great apes from imminent extinction and
a feast or potlatch; this attitude should again be
to
formulate at least a middle-range conservation
enthusiastically embraced.
scheme
As
by implementing
GRASP may
scheme,
country
measures
to
be considered a middle-range
mission should be to help each range
its
formulate a national great ape survival
to
'Progress' appears to be currently regarded
save them.
as the sole ethical purpose
However,
human
During most of
network. Limiting road construction and logging,
more
along with the introduction of Forest Stewardship
conservatism or respect
management, are per-
haps the most important elements

I
consider
short-term and
with
conservation measures, and so
middle-range
will confine
myself
taking a long-term perspective for great ape
dominant
Most
am
live in Africa,
is
and
how many chimpanzees
answer
Upon hearing
"Oh! There are
still
so
that
this,
still
perhaps there are

the usual response
many chimps!" People just
do not stop to consider that even the smallest

satellite cities of
Tokyo or Osaka contain more than
100 000 humans.
we do
We
are so anthropocentric that
not think twice about the fact that
we
overpopulating the Earth at the cost of other
are
living
existence.
that
history,
and even
nowamong
communities around the world,
of
often
customs remains the
this rather than that.
responded, "Oh, because
of the time, they

to
my Tanzanian
asked
why they were doing

do
this."
We
my
rarely recognize
most modern developments provide only short-
term benefits by wasting materials.
Although
often asked
only 100 000.
ethic.
grandfather used
conservation.
I
traditional
assistants
feel that the three chapters that follow deal
excellently
to
to
human
Europe after the 18th century Enlightenment.
plan and to integrate these plans into a global
Council principles of forest
of
became widely accepted across
this only
the
anthropocentrism
human
race,
the decrease
many
is
is
ultimate
result
of
today's
the endless expansion of the
believe that the primary cause for

in
the
number
of great apes,
and
other species of wild fauna and flora besides,
the ever increasing
industrialized
demand,
particularly
in
the
world, for raw materials: cheap
timber, cheap agricultural products, cheap natural
resources. So, perhaps those of us

trialized
world should
first
in
the indus-
be asking questions of
215
World Atlas
ourselves about our current consumerlst attitudes
and
tiabits.
witti
any Integrity
is
Only then, for example, can

witti
we work
those range states where
it
practiced to discourage the eating of great apes.
There are other ways forward,

years now,
have been engaged
too.
For four
the Great
in
some
at least
of
some
Accordingly,
tected
nominated based on
Internationally.
universal value, encap-
This
is
one
of
the
guiding
reasons
to
propose
is
Although ecotourism and even research have
long-term
scientific
locally based, while organized
Gombe, Mahale, Karlsoke, Wamba,
Kahuzi-Biega, Bossou,
Budongo,
Tal, Kibale,
and others each have such a team

Kallnzu, Bwlndl, and
granting World Heritage Status to the great apes.
be invaluable.
to
would suggest that any pro-
area should have a
research team that
its
Although
survived the crises, with the support
determined scientists proving
believe that a World Heritage Species should be
sulating scientific, cultural, and conservation ideals.
to their assistants.
the study populations suffered devastating losses,
Ape
World Heritage Species Project (GAWHSPI as
scientists continued to visit their study sites
and provide salaries
long-term club. The
Kutai,
of scientists.
Moukalaba may soon
join this
could be extended to
list
all
occasionally had negative effects on the health of
the protected areas containing great apes, possibly
great apes through the introduction of anthropo-
with international nongovernmental conservation
zoonotic diseases, large
numbers
of
apes have been
protected by park rangers funded at least partially

by ecotourism,
for
example
in
Mahale, United
organizations involved
for
their 'own' animals,
to play.
John Gates
ttie
in
his excellent
research fees can also be encouraged, with range
trust funds to
term research.
some
who
of
perhaps we - as proposed by
book Myth and Reality
Rain Forest- should be encouraged
states regarding these as one source of long-term
finance for protected areas, as has been the case
helping to provide funds
monitor great apes every day are very protective
Republic of Tanzania. Thus, ecotourism has a part
Both academic and conservation-oriented
in
long-term research. And as researchers
ensure the
viability of
in
to establish
continuing long-
In
East African countries. Although research
fees might be lower than tourist prices, researchers
should accept
some
costs, since their
work provides
individual benefits to the researchers. Short-term
success
in
the conservation of great apes In the last
20 years has been disrupted by war and

conflict.
216
armed
However, during even the worst period.
Tosliisada Nishida
Executive Director, Japan
Monkey Centre
Challenges to great ape survival
Chapter 13
Challenges
to
great ape survival

Lera Miles, Julian Caldecott.
AND Christian Nellemann
Great apes
are endangered because people
Threat status classification
are bringing into their world both deliber-
The Red
ate change, such as land clearance,
World Conservation Union
change, such as forest
accidental
fires.
and
This
destruction of tropical moist forests puts countless
numbers
of
species at risk
of extinction.
Hence, the
challenge of great ape conservation cannot
management and
disentangled from the
tropical forests as a whole. That said, great
particularly vulnerable
because
it
is
be
List of
Threatened Species of lUCN-The
which species are

servation
action.
losses are an important criterion for the
more over
whichever
is
classified
as Critically Endangered;
ten
the longer), or
them, they reproduce slowly and
percent
many human
ape species are
diseases. The main
it
is
classified as
in
Endangered categories
and fragmentation due
to logging
ticular
agriculture (particularly
in
hunting (particularly
in
for
West Africa and Southeast

Southeast Asia]; and
West and Central
Africa).
Potential sources of further risk include diseases,
human
conflict
and mineral extraction. Demand from
overseas consumers for luxury resources such as

tropical
timber and cheap staples such as palm
contributes to
and intensity
many
of these pressures.
of threats
The range
have led many observers
conclude that great ape numbers
oil
will
Red
List.
If
is
expected
Endangered.
to,
if
then
it
is
by 50-80
All the great
the Endangered or Critically
threats to their survival are habitat loss, degradation,
and clearance
con-
years (or three generations,
or
apes are
easy and often
of
has declined by 80 percent
future of
profitable to shoot
a guide to determining
Recent or expected population
a species' population
are susceptible to
Asia); forest fires (especially
is
most urgent need
in
of the
Red
List.^^
Of par-
concern are the Sumatran orangutan, the
mountain
gorilla,
and the Cross River
which are classed as

Table 13.1). The
Critically
gorilla, all of
Endangered (see
Red /./sf coding system
also reflects
Vu Danh Viet/UNEP/Topharr,
Forest fires are one

of
many threats
to
ecosystems, particularly
in
Southeast Asia.
to
further
decrease, and rapidly, within 10-20 ygars.'^"-'"'"'-'^^
PREDISPOSITIONS TO ENDANGERMENT
Life history
and vulnerability
Great apes have relatively low reproductive rates,

long lifetimes, and long 'childhoods' (Table 13.1).
This combination of factors
makes
their populations
very vulnerable to high rates of adult mortality, from
which they cannot easily recover This, combined

with the requirement for a large area of natural
habitat,
is
the ecological factor at the root of the
vulnerability of
apes
to the
impacts
of
humans.
217
World Atlas
the type of evidence on which the classification
1.1.1.1 (habitat loss
has been based." The Sumatran orangutan, for
example,
3 (harvesting/hunting).
falls
into
category
the
Endangered (CRI A2bcd'. These

bers
mean
'Critically
letters
.3.3 (habitat loss
through shifting agriculture);
through wood extraction);
and num-
The lUCN system therefore condenses expert
that:
opinion and scientific evidence about the status of a
the
orangutan
A (population
criterion
fits
species, subspecies, or population. There are 352
Endangered and 162
decline);
of type 2 (reduction of
over the last 10 years or three generations];
based on evidence
types
of
abundance), c (continuing decline

habitat'),
and d
(index
Critically
mal species on the 2004 Red
common
this Is a
endangerment
of
warning
in 'quality of
tale of
threatened extinction." The
apes can be seen as an early
of
of the loss of
many
beings and the great apes
The major threats are also categorized.

Sumatran orangutans, these
Table 13.1 Great ape reproductive characteristics and

Taxon
Chimpanzee [Pan
troglodytes]
Western chimpanzee
(P.
t.
For
own
(,
to
our
kin.
Minimum
Reproductive
Maximum
Estimated no. of
Year of
age at first
pregnancy
Interval In
lifespan
Individuals
estimate
adult females
10-13
A.4-6''
40-50"
172 700-299 700^'
2003'
EN A3cd
21000-56 000"
2003"
ENAlcd+2cd
76 400-119 600"
2003"
EN A3cd
70 000-116 500"
2003"
EN A3cd
2003"
ENAlcd+2cd
2001
EN A2cd
verus]
IP.
f.
troglodytes]
t.
human
their case par-
lUCN
Status'
'"
schweinfurthii]
Central chimpanzee
[P.
makes
status
Eastern chimpanzee
(P.
species that are less
resonant for us: for these are threats
ticularly
are:
/?ec//./st
reminder that
well known. The close relationship between
(level of exploitation).
mam-
Endangered
List, a
000-8 000"
vellerosus]
13-15"
Bonobo [Pan paniscus]
L&'''
50-55'"
10 000-50 000
(to
Western
gorilla [Gorilla gorilla]
Cross River
4-6"
j-c6i. 160
'''"'"
94 500-1 10 000'
2000"
EN A2cd
250-280"
2004
CRA2c;C2a(il
94 500-110 000
2000
EN A2cd
gorilla [G. g. diehli]
Western lowland
gorilla [G. g. gorilla]
3.9-4.6"'"
Eastern gorilla [Gorilla benngei]
Mountain
35-45""'"'
gorilla IG.
b.
IG. b.
2005
i6l.8i, 101. 102
3.9"
benngei]
> 100 0001'"'
700'
?f.g 12'
4.6"'
graueri]
EN A2cd
2003
CR
2005
EN A2cd+
2004
EN A2cd
2004
Not assessed
2004
EN A2cd
C2alil
3cd+4cd
Bornean orangutan [Pongo pygmaeus]

Northeast Bornean orangutan
IP. p.
8'"
ll-lS'*
45"
45 000-69 000"'
mono]
11
Northwest Bornean orangutan

IP. p. pygmaeus]
Central Bornean orangutan IP
Sumatran orangutan [Pongo
p.
000-21 000'
640-3
>40
wurmbii]
260''- '"
500'2'
7 334'^'
abelii]
a EN, Endangered: CR. Critically Endangered; for a
full
explanation of tfireat criteria see
lUCN
(20001
lUCN Red
List
2004
EN A2cd
2004
CR A2bcd
Categories and Criteria Version 3.1
finp://www.]ucn,orgAfiemes/ssc/redlists/RLcat52001booklet.html. Accessed Marcfi 28 200^.

b Cfiimpanzee and western gorilla estimates were collated by
Thomas
Butynski [2001 ",
the country profiles for furtfier details.

c
Based on eastern
Based on western lowland
gorilla,
gorilla.
e Recent decline due to Ebola hemorrhagic fever not quantified.

f
No
Reduced from
218
data; fieldwork
was being undertaken
17 000 8 000
in
1998.^^
in
2005
to
estimate the extent of the decline,
2003^''|
and include estimates ranging from 198A
to
2003. Please consult
Risk and uncertainty
As our ecological knowledge has increased since
Table 13.2 Estimates of Bornean orangutan populations through time'
the mid-20th century, our estimates of the popu-
apes have also increased, even
lation sizes of great
Year
though their populations actually declined during
19605
The Bornean orangutan
1970s
period.
this
example Isee Table
This
13.21.
a typical
is
because early
is
estimates did not always recognize the breadth
of
habitats and regions occupied by the great apes,
and further errors have arisen from the variety
of
Estimate
Trend
450-900'"
declining
250', 2
1990
000-3 000'
declining
30 000-50 000''
not staled
15 953-24
1995-1996
declining
>3 500'"
19805
497'""
declining
45 000-69 000'"
200/1
declining
sampling and extrapolation methods used. These
and ape nests are not
often rely on nest density,

easily visible
in
the forest: radically different results
may be obtained from surveys on
or
foot
See also Table 10
Sarawak only
Sabah
4.
only.
by
helicopter, for example.
Nevertheless,
we can be
confident that the
species with the smallest populations are the eastern gorilla and Sumatran orangutan
that the scarcest subspecies
13.1),
River gorilla, and that the
the Cross
is
chimpanzee
is
many chimpanzees
times as
gorillas, four
Bornean orangutans or bonobos, and about
as
Taxon
Year
Percent decline
Red List'
Trend
over period
the most
numerous great ape species. There are, very

roughly, at least twice as many chimpanzees as
western
Table 13.3 Sumatran orangutan decline, based on 2004 lUCN
(see Table
1992-1999
Pongo
abelii
lib
declining"
1992-2000
Pongo
abelii
>50
declining"
2000
Pongo
abelii
17
declining"
See also Table
,3
many chimpanzees as Sumatran

may be over 90 000 each of the
30 times as
orangutans. There
The only other
central and eastern chimpanzee.
numbers
this
is
exist
similar
in
the western lowland gorilla. Contrast
human
with the global
which was estimated

of
may
subspecies that
great ape
population
2005,
in
at 6.^65 billion: the equivalent
about 27 000 people for every chimpanzee.
'^^
in
some
provides
(see
great ape numbers. The 200^
Table
Red
of
List
estimates for Sumatran orangutans
13.31.
also
It
states
Bornean
that
orangutans, chimpanzees, and eastern gorillas are

declining
in
numbers, but that there
is
insufficient
information to judge the trend for the western

gorilla
and bonobo." The dual impacts
and bushmeat hunting
western
gorilla
in
the heartlands of the
and chimpanzee range
Basin are unquantified and
may
reduced
both
populations
of
Ebola
of
in
the
Congo
already have
these
depend
on
much
'common'
build nests in trees
food
sources
mountain gorillas forage mostly
at
and
(although
ground
level
within a herbaceous 'canopy']. Even chimpanzees,
the
species most likely to be found
which
to
sleep and feed.
survival of the great
apes
fate of the forests in their

It
IS
to
in
open
forest in
Hence, the long-term
will
be determined by the
range countries.
maps
hard to find global
of deforestation.
The Food and Agriculture Organization
of the United
Nations (FAQ) collates information about forest area

by country, including estimates of the
amount
of
change. Forests are defined by FAO as areas with
canopy cover
of
more than
10 percent, including
both natural and plantation systems. Estimates of
change
in
natural forest are not provided separately
from estimates
of
change
in
forest cover, but this
information can be obtained approximately by sub-
species.
tracting the
HABITAT LOSS
AND FRAGMENTATION
The great apes are creatures

forests,
apes
arboreal
woodland or farmland, need access
There are few figures on the actual rate

decline
of the time, all the
although not
all
of
tropical
change
in
plantation forests from the
total (see Table 13.4).
moist
are equally arboreal.
Deforestation
is
development decision
sometimes the outcome

(e.g.
Orangutans are seldom seen on the ground,
forest
whereas the more
resettlement scheme), but
relatively little
time
in
terrestrial
bonobos spend
the trees. Nevertheless,
most
with
sequence
of
oil
palms,
or peat
many small
of a
replacing lowland rain
it
forests
with
can also be the con-
actions that coalesce
219
World Atlas
(e.g.
settlers
and
and their smallholdings multiplying along
a forest
whole
hillsides
moving
on,
create a deforested environment
to
highway).
when one form
can also occur
It
turbance (such as logging! allows the forest

out enough for
happened
burn
to
it
Borneo
In
fires
if
to dry
are set nearby (as
1997-1998 and
in
Sumatra
Deforestation can be irreversible
2004!.
in
In
of dis-
if,
for
example, exposed soils are badly leached and eroded

or
fire-adapted
if
of forest.
factor
in
Southeast Asian
the
orangutans
especially
live,
human
soils are scarce,
in
important
forests
Borneo. Here
where
fertile
populations sparse, and
land-use practices are highly consumptive
of
space
few
Every
natural forest cover
range states, 1990-2000"
in
Change
Natural forest
2000 (km'l
Angola
Annual change
(percent of
1990-2000 (km'
1990 figures!
-12 630
-0,18
696 150
Burundi
In
natural forest
211
season
of forest,
279
-8.58
-22 820
-0.88
-1
237 780
229 027
-2
983
-0.13
Congo
219 767
-2 213
-0.10
Cote
the
69 327
-27 703
-2.86
351 103
-53 837
-0.38
030
-0.56
d'lvoire
Democratic Republic
of
Congo
cut
in
In
17 520
-1
Gabon
217 896
164
-0.05
62 590
-12 230
-1.64
Guinea
69 042
-3 678
-0.51
-1
Indonesia
21855
-2 165
-0.90
-168 695
-1.51
33 625
-8 725
-2.06
935
-0.70
Mall
131 715
Malaysia
175 425
-40 375
-1.87
Nigeria
128 239
-45 261
-2.61
Rwanda
462
-3 228
-875
Senegal
59 418
-6012
-0.92
Sierra Leone
-9
especially the case
is
The
In
filled in after
use, and continue to
lower the local water table

Ivlajor
ENSO-llnked
1982-1983, 1991, 1994, and 1997-
997-1 998
were so widespread
fires
pollution,^
and
occurred
2002 and 2004.
in
many
such
other
that
areas,
fire
haze
events
also
and farming combine
to
create a stable patchwork landscape, often with
moist forest retained
In
gullies
and
valleys,
maintained grassland on the slopes

Ivjuch
of
in
and
fire-
between.
the habitat of the Cross River gorilla
like this, for
Nigeria.
example on the Obudu Plateau
in
Division of the Cross River National
Small populations can
find
is
the
Park
themselves
disperse to other or larger areas. Corridors
of
farm development or housing along roads can

also fragment a forest landscape, and with
ape population.
Some
of the
measures
it
a great
that con-
servationists use are designed to offset this effect,

for
example, by linking habitat blocks with forest
Fragmentation
corridors.
reduces the size
of the
population, thus reducing
and increasing
its
is
hazard because
It
gene pool within a breeding

Its
genetic heterogeneity
vulnerability to the effects of
10 490
-3 610
-2.56
609 865
-100 265
-1.41
vulnerable
386 761
-8 939
-0.23
as disease and forest fires."
472
-9 358
-1.84
41
where canals are
forests to float out logs, since
swamps and
occurred
998.'
and the
opening and allowing the drying
Sudan
Uganda
Southern
onset of the
trapped within such patches of forest, and unable
951 155
Liberia
to the
millions of people wer-; affected by atmospheric
to
Guinea-Bissau
in
long after logging has ended.
In
Ghana
Nino
El
Southeast Asia. Together with the
peat-swamp
Okwangwo
Equatorial Guinea
the
major
of
schemes.
these droughts exacerbate the spread of
This
fires.
In
Cameroon
and the location
years,
effects of logging
fires
Year
Borneo as more-or-
local tradition of using fire to clear land,
drain the
in
of
(ENSO! causes a delay
they are seldom
Table 13.4 Decline
large areas
In
infrastructure projects or resettlement
rainy
particularly
interior
permits,
plantation
are then maintained by regular burning.

Is
same time hunt
less 'empty' land, for the allocation of logging or
Oscillation
fire
one or two harvests before
tor
at the
Meanwhile, central government planners
[Imperata cylindrica], become well established and
Large-scale
and
treat the
often
such as alang-alang
grasses,
Local farmers are used to clearing
forests.
inbreeding. Isolated populations are also potentially

to
catastrophic or
random events such
HABITAT DAMAGE
a These estimates of deforestation between 1990 and 2000 were determined from
Resource Assessment IFRAj data by subtracting

to
FRA
2000."'
'5
from
tfie
cfiange
in all
tfie
(crest area',
FAO Forest
'cfiange in plantation area',
FRA
2000.'^
FRA 1990
The
last
half of the
establishment
ber and the
20th century
of a global
market
availability of capital
saw both
the
for tropical tim-
and equipment
within tropical countries to enable widespread
220
Box
LUCKY GORILUS?
13.1
Large animals that reproduce slowly are usually the

be threatened with extinction. Gorillas are
first to
the largest primates, and only the other great apes
reproduce as slowly as they do.
Nevertheless,
neither the western nor the eastern gorilla
as one
Table
13.11.
is
listed
most threatened primate species
the
of
species recognized
in
the
(see
the
296 primate
lUCN Red
List are listed
Over 6 percent
of
as Critically Endangered," but
out of six great
five
Ian
Redmond/UNESCO
lucky gorilla?
ape species are only Endangered, lOn the other

hand, one subspecies from each
species
Why
is
among
Whether
determined
the
most threatened
a taxon
to
danger
primate
of extinction
Red
be blessed with
by
biology.
its
some
luck
in
Gorillas
both areas.
(II
one
of
of the less
on herbaceous food as opposed
any
In
food exists
in
quality food.
great apes
ing
it,
can survive largely

to relying
higher energy content, such as
meat.'"
on food
fruit
or
forest, this relatively poor-quality
Luck with threat
fruit is in
short supply, the other
must expend considerable energy seek-
while gorillas start to eat
more
low-quality
food. Despite their large size for a given
can therefore survive
of food, gorillas
swamp
in
gorillas
forest,
in
in
to
them
is
West African
of gorillas
from logging or the bushmeat
human
population den-
than they or other animals do elsewhere.
Luck with threat

Not
swamp
all
human
(3)
parts of the range of gorillas have a low
population density
Some
of
smaller
gorilla in eastern Central Africa.'^
and rebellion
can occur
at high densities
as biologists have only recently
forest
remain
occur
in
this
in
mass slaughter
the highest den-
the range of the eastern
The chaos
of gorillas,
such as
affected by logging
same time some
protected areas
in
war
that of the
of the very best
Africa occur within the eastern
through
gorilla's range;
and genocide, the Virunga mountain
in
swamp
forest,
and associated changes arising
from drainage and subsequent
Park.'" At the
intact, particularly in
Southeast Asia these have been greatly
ulation
pop-
gorilla
has flourished."^
(western lowland and mountain gorillas]
many
seem
less
other primate species

is
we
safe.
Ebola hemorrhagic fever epidemics appear to have
11)
Mammals and birds are more often threatened

where human density is higher,^' but very few humans live in the vast areas of West African swamp
forest.
low human density and large geographic
If
range both make taxa safer, there are grounds
war
the vicissitudes of
Of course, just because two gorilla subspecies
threatened than
fire.
all
cannot assume that the genus as a whole
Luck with threat
of
has occasionally led
area
gorilla's range, vast
Congo." Orangutans also do well

in
gorillas
eastern lowland gorillas of Kahuzi-Biega National
discovered."'"'" Within the
but
forests cur-
perhaps now the majority
forest,
Africa, suffer less
sities
swamp
sities in Africa
12)
Western lowland
areas of
The
difficult.
trade under comparable
to
swamp
extremely
the
of
few people, but access
amount
areas of natural habitat than can the other apes.
in
(2)
Not only do the interior
greater abundance than does high-
When
Lucky biology
emerges as
threatened primates; over 50 percent
other primate species are more threatened.'"
still
Gorillas, unlike other great apes,
with a
When ranked
using these criteria, the western gorilla
rently host very
Lucky biology
be less threatened
List suggests.
by the nature and intensity of
first
the threats, and second
appear
is in
of
may
think that western gorillas
than the lUCN
are the gorillas, the largest of the pri-
mates, not
species'?
the gorilla
of
Endangered.)
Critically
IS
to
eliminated western
lowland gorillas from
large
areas of eastern Gabon. '^' Despite decades of conservation effort, the threats to gorillas from land-
use changes and hunting are
still
very real.
Alexander
H.
Harcourt
221
World Atlas
officials,
to
the impact on the forest ecosystem
How
be severe.
severe
is
likely
depend on various
w/ill
including the intensity of the operation.
factors,
Southeast Asian dipterocarp forests are often
much
harvested at a
characteristic
higher rate than has been
similar operations
of
African
in
because the dipterocarps grow so densely
forests,
and are such valuable timber trees.

Studies of timber-producing tropical countries
have revealed a consistent pattern. Government

forest sen/ices have tended to function reasonably
well until a combination of
markets made
it
possible to
new technologies and

make huge profits from
logging.'^' At that point, multinational
were often
country
companies
by the government
invited
in
each
harvest the forests on a profit-sharing
to
basis. This generally required both that the gov-
This orangutan
ernment's own forest service be
in
Central Kalimantan
so that
is
it
partially disabled
could no longer insist on sustainability,
and that legislation protecting the forests be
showing signs of
starvation as a result of
rewritten to allow long-term public interests to be
habitat disturbance.
overridden by short-term, often personal, financial
occurred
interests. This
industrial-scale logging.
Even the best-managed
mechanized logging has
a strong
local
impact on the
environment. Roads and log-pounds are
built
soil;
trees
must be
felled,
crushing their
Conversion of great ape
neighbors, and then dragged around, damaging
habitat to agricultural
other trees. Bearing
in
mind that
this
is
being done
a tropical, high-rainfall environment, often on
land. This area lies
in
between Bwindi and
steep terrain, and frequently far from the super-
Mgahinga
in
Uganda.
vision
of
professional
foresters
and
forestry
Gordon Miller/IRF
different
in
in
Malaysian Borneo, and during the 1980s and 1990s

in
Indonesia.
using heavy equipment, compacting and exposing
the
times
at various
example during the 1970s and 1980s
places, for
Indonesia
still
held most of
forests as late
its
as 1950, but over the following 50 years forest

cover declined from
The
620 000 km'
rate of forest loss
is
still
lowland forests being most at

lowland forests
will
to
980 000 km'."
accelerating, with
risk.
At current rates,
disappear entirely from Sumatra
and Kalimantan (Indonesian Borneo! within 10-20
and
years
perhaps even
sooner.
Indonesian forests are allocated
to
ivjost
export-oriented
logging concessions. Barito Pacific Timber Group,
the
holder
largest
of
logging
concessions
Indonesia, exported over 9^ percent of

tion in
is
2001." Extensive
its
in
produc-
illegal logging for
export
also a major problem within the industry.""
program funded by the UK government found that

73 percent of
all
logs
in
Indonesia
documented sources,'" while

logs have for years provided
came from un-
illegally
exported
raw material
for the
Malaysian sawmill industry, which has far too great

a capacity for
its
own
(legal!
in-country supplies.
The Indonesian government, supported by the

European Union, United States, and others, has
been trying
over
this,
for several years to gain
and
in
some
control
July 200i proposed introducing
the death penalty for those found guilty of illegal

logging."
222
Box 13.2 SATELLITE ANALYSIS OF THREATS TO
decades,
GOMBE CHIMPANZEES
Research Center have been tracking chimpanzees
members
staff
As so much
to collect data daily
On January
chimpanzee, flew
Ham's
missions.
pave the way
Ham,
1961,
31
one
in
flight
a four year old
success that helped
manned space
for the United States'
program.'" Space technologies
flight
NASA space
of the first
was
male
now have
each
unique resource
the behavioral ecology and conservation needs of

their species.
Gombe's chimpanzees face many threats
the
The park
km^
and are
save chimpanzees and other endangered great
chimpanzee numbers are declining as a
apes from
disease, poaching, and habitat loss.
extinction.
we need
the great apes are to survive,
measure
success
objectively the
action. This
means we need
to
conservation
of
extensive information
on habitats and land-use patterns that include suf-
the great ape ranges occur
where
the world
of satellite-based
now make
different characteristics
map
information
little field
multitude
developing parts of
in
is
available.
sensors with
possible to
it
the location, extent, and magnitude of certain
types of
human
Satellite
imagery has been used
understanding
great ape ranges.
activity within
is
established,
of the threats to
support conservation efforts
Gombe
Gombe,
an excellent
be possible
employed
to
improve the
to
threats to
mosaic
of
to
be
chimpanzee
Gombe chimp-
evergreen and deci-
duous forests and woodlands, so
imaging
satellite
evaluate the extent to which these
habitats have been lost from the
Landsat
good
reduce or eliminate
to
habitats at different spatial scales.
to
imagery has proved
m mapping
anzees depend on
has been used
Once
monitor conservation success.
satellite
tool
area;
in
result of
Gombe
region.
have been orbiting the
satellites
to
Earth since 1972, continuously collecting images of
National
the Earth's surface. The accumulated data archive
chimpanzees and
in
may
it
these threats, and

At
only 35
is
these threats and their causes
of
practical strategies
improve our
to
in crisis.
understanding
both time and space. Unfortunately,
ficient detail in
are a
seeking better understanding of
in
potential to contribute critical information to help
If
includes
Park, United Republic of Tanzania.
Gombe, Jane GoodaU's pioneering work and
multiple
images
of
Gombe, permitting
images from
comparison
of satellite
groundbreaking discoveries about chimpanzee
'Vegetation
difference indexing', which quantifies
behavior helped to narrow our view
changes
In
the gap
of
between human and nonhuman beings, teaching us

a great deal
about our own place
on the shores
Tanzania.
in
Gombe
National Park
1968 and was the
to protect
in
nature. Located
Lake Tanganyika
of
western
in
was
established
park created specifically
first
Gombe
chimpanzees.
hosts one
of
the
world's most longstanding sites for the study of
animal behavior Research began
Until the 1980s,
in
1960;
timber from West and Central
was considered
value,
which limited the pressure posed by the
selective
logging that
be
to
was
of
low commercial
taking place. All this
in
1972 and 1999. This revealed forest destruction
Gabon's forests were
of
allocated as logging concessions;" and log pro-
duction
mVyear.
had increased
Meanwhile,
in
to
some
2.5-2.7
palm
miombo
of loss of local tree cover
million
Cameroon, more than
70 000 km^ (76 percent! of the country's forests had
either been logged or allocated for logging con-
plantations, along with

(dry deciduous!
wood-
occurred outside the na-
on village lands as well as within forest
tional park,
continued overleaf
cessions, and satellite images have revealed that
networks
of
new
logging roads had spread into what
had been considered the least accessible forests
in
the country.'"' Extensive logging had also occurred

in
half
of
lands for farmland and charcoal. The vast majority
the
more than
to oil
massive clearing
Southeast Asian forests became depleted.

2000,
green vegetation cover, was performed
in
and conversion
changed dramatically during the 1990s, as the

By
different dates.
using satellite images captured during dry seasons
for
Africa
known about
is
Gombe chimpanzees
the
individual,
Gombe Stream
the
of
the
Rio
Muni area
DRC and Congo
Forest,
and
range.*'-
in
of
Equatorial Guinea,
in
Mayombe
sections of the
other parts of the western gorilla's
''='"
Many
of
the timber concessions
been awarded within the range

According
of
DRC have
in
the
bonobo.
some 2A
now under
to the national forest service,
percent of the range of the bonobo
is
223
World Atlas
Figure
A Change
in
Gombe
forest cover in the
expressed
terms
is
reserves,
in
areas
known
be inhabited by chimpanzees (Figure
to
ttiat
in
of
region, 1972-1999
the Normalized Difference Vegetation Index INDVII.
Forest cover
are less well protected but
The analysis also revealed an increase
Gombe
forest cover both inside
within
patches
protected
Kitwe (Figure
the Tanganyika
of
forests
such as
Institute.
demonstrates the enormous potential
This
restoration of forests and
Kigoma
region.
If
large
for
miombo woodlands
enough patches
of
in
these
chimpanzee population
products
now account
trade recorded
d'lvoire,
Many
in
for
(e.g.
Alpi],
of the
logging companies operating
such as Denmark
in
human
cise satellite imagery has recently
able that can detect objects
Logging
and Germany
(e.g.
(e.g.
Danzer, Feldmeyer, and
in
map
Most
western
in
More pre-
become
the 0.6-4
tropical moist forests
in
is
avail-
scale
complex
ecosystems, having wide-ranging effects on the

various species present.""'
(t
is
not always clear that
logging at moderate intensities, considered alone,
wholly negative impact on great apes, which
robust and
Italy
length.
process that interacts with varied and diverse
Group), France
in
therefore difficult or impossible to map.
has
DLH
Gombe
for the
settlements, roads, and farms
are
(e.g.
hope
Tanzanian landscapes are smaller than that and
the
in
is
the long term.
Historical Landsat satellites can only
European Union coun-
Rougier, Thanry, and InterwoodI,
Wonnemannl."
IIU
10 percent of
Equatorial Guinea, Gabon, and Liberia.'"
Congo Basin are based

tries,
more than
Cameroon, CAR, Congo, Cote
in
land features larger than 57 meters
logging concession;'" other observers calculate
the figure to be as high as 55 percent."" Forest
all
strategically important
in
forest remnants, then there
at
Catchment, Reforestation, and
Education Project of the Jane Goodall
the
in
National Park and
which have been restored by
A],
habitats can be restored
places for chimpanzees and connected to existing
A|.
dietary needs.
duces
fruit
mobile and have unspecialized
Where
the process of logging re-
availability,
however, then carrying
capacity at least for the orangutan will inevitably

decline.
The more folivorous and
terrestrial gorillas
Figure B
Fifi's
This 1-meter
progress (yellow dots! through
IKONOS image was
The effectiveness
of
mapping heterogeneous
collected
new technologies
farms, settlements, and paths
strated
in
(a
demon-
is
-meter IKONOS imagel. The
yellow dots represent the location of
Fifi,
a female
chimpanzee, as she was being followed

minute intervals
which many
1999. This
in
human
activities
is
at
15
the fine scale at
Impact directly on
chimpanzees, so such images are extremely useful

In
is
draped over a
Greater
digital elevation
accuracy
and
higher-resolution
imaging costs more. The dozens

by various satellites planned
increase the range
of
in
model.
sensors carried
of
the near future will
options open to researchers.
The cost-effectiveness
of
these tools depends on
matching technological advances with high-quality
and the integration
questions,
obtained
Improved
into
the
of
efforts
results
great
in
ape
conservation.
planning, implementing, and evaluating conser-
vation
measures
in
the
Gombe
Lilian Pintea
region.
and the more adaptable chimpanzees are
likely to
be less affected, even by moderate logging.

are
National Park
in
tree canopies along with
human
Figure B
Gombe
July 2000 and
in
If
they
from their normal home ranges,
displaced
however, this can cause stress and disruption

social
interactions with
to
neighboring groups and
communities.'"
What
is
may
who
chainsaws
of
much
drive the bulldozers
who
roads,
is
that
and wield the
logging operation, their
followers, and people
logging
clearer, however,
camp
arrive later along newly
are
likely
to
want
to
their food supply through hunting.
They
also wish to supplement their finances by
selling forest products,
if
great apes as food, and
Many will regard

smoked ape meat as a
they can.
commodity; these additional hunting pressures are

a
very
the workers
opened
supplement
serious threat associated with logging. Once
Industrial logging
has run
its
course, moreover,
plantations, farms, ranches, fires,
and
alien invasive
species tend to enter the logged areas, creating a
new ecosystem
canopy
forest,
with few patches of native, closed-
and therefore few or no great apes.
225
World Atlas
bonobos, chimpanzees, and even gorillas as
infant
and
pets, for zoos,
hundreds
for private collections;
chimpanzees were exported
of
for
biomedical
research, a trade that continued into the 1980s on

existing
permits,
despite
under the
restrictions
Convention on International Trade
in
Endangered
Species of Wild Fauna and Flora (CITES).'" The

profitability of this illegal trade continues.
Apes as food
Consumption
many
ape meat
of
apes as objects
primates
is
provided
some
both
of
for sale at a
Kasese market
stall,
HUNTING AND THE BUSHMEAT TRADE

Historical
protection
Borneo and
of
the Congo.
including
uses,
traditional
the production
of
charms'. Gorilla bones are an important ingredient

of
much sought
amulets, which are
protective
healers." Chimpanzee and gorilla
after by local
body parts are important elements

medicines
West Africa'" and
in
Sumatra, the
staffs
and wands
Congo.""
in
of
traditional
in
In
shamans were
often decorated with orangutan hair''' In Borneo,
where head-hunting
traditions
both wide-
are
spread and recent, orangutan skulls can fetch up
US$70 m towns
to
Kalimantan, apparently as a
in
human
less illegal alternative to using
skulls (see
also Box 13.3).

in
apes as pets and research
in
There was a booming
in
inter-
orangutans between the 1930s and
them became
1960s, although hunting
192i
hunters with
provided
historically
additional income.
national trade
hunted
Sumatra and
in
1931
in
illegal
Sarawak.'"
in
the
In
for
bushmeat, which
into a profitable activity Isee
]3.A].
great apes
been a harvestable resource
for subsistence, or to
modern indigenous peoples
have
to
be
meet market demands
meat and medicinal products.
for
Prehistoric and
of Indonesia are
known
to
have hunted orangutans extensively for meat and
to
have preferred ape meat
to
game.''' The Batak people of
many
other kinds of
Sumatra
believe that
make them strong, and this

belief has prevailed into modern times. Throughout
Central and West Africa some communities have
eating the
meat
will
traditionally eaten
market demand
dish
fit
chief
powerful
for
who
ape meat and there
for it," despite
prohibitions. Gorilla
The trade
animals has
traditionally
parts
in
population
however, as has the
demand
some communities,
For
for a variety
Human
exchanges have altered
cultural
increasing market
Boxes 13.3 and
Apes have long been valued as sources
Sumatra and
in
Africa.
traditions,
has turned hunting
markets
Manlena province,
reverence, or as too similar
of
forbidden under Islamic law, which has
movements and
some of these
Bushmeat
in
themselves, or as unhealthy to eat."* Eating
to
Pierre Kakule Vwtrasihikya
limited by tradition
is
sometimes because people regard
places,
meat
men
in
not serve
did
particular
some
in
it
is
a high
customary or legal
is
seen as a
cultures, so that a
to visiting
dignitaries
risk
embarrassment. Gorillas are eaten by
some who
believe they will thereby gain strength;
would
and chimpanzees are eaten

their luck
in
the hope of acquiring
and cunning.'"'
1980s, popular movies and television soap operas
created
new demand
especially
in
for
orangutans may have been imported

the
pet
trade
orangutans,
pet
Far East. An estimated
the
between
price of an individual ape
1995
in
into
and
000
Taiwan
1999;
for
the
the private market
was reportedly anything between US$11 000 and

US$20 000. Much of this trade has now been
stopped, but
it
is
highly profitable
where
it
does
occur There has also been a lucrative trade
226
in
Scale of bushmeat trade
By 2000, the
to
be worth
illegal
bushmeat trade was estimated
nearly US$1
billion
annually," a
proportion of which involves great ape meat."'*'^^'

'"
It
has been
facilitated
by the opening up of
formerly remote areas by logging companies,

especially
in
Central Africa,* "* '" where a single
chimpanzee or
gorilla
equivalent of US$20-25.'
carcass
can
fetch
the
Box 13.3
HUMAN
BELIEFS
AND TRADITIONS
These
traditional attitudes reflect the special
nature of apes, rather than their protection status.
The cosmopolitan,
'western' culture
rationalist,
is
not the only one with strong opinions about apes.
The ethnographic record shows
that traditions
and
beliefs
about apes are widespread, with both
positive
and adverse
results. In
Borneo,
many
Iban
believe that after death an ancestor
becomes
member
and
species
of a particular
[tua]
in
this
There are
In
where the
great ape
such
a tua species."
Other peoples
of
Sumatra and
Borneo saw orangutans as representatives
war
of their
human rulers who had fallen

humans who refused to speak for
god, as former
from grace, as
fear of being enslaved, or as their forest cousins." In
the northern inner

a
fallen brother'
again.
Congo
who
Wehdjeh clan
of the
Sapo
people consider themselves to be relatives of the
chimpanzee, from
skills
is
whom
their
derived,'" while the
knowledge
of forest
people of Congo
Vili
hold that chimpanzees are reincarnations of people.

Similarly, people in the
believe that
of
chimpanzees shelter the
elders, so killing
many
Boe region
of the
them
is
taboo.
In
Guinea-Bissau
Cote
totem and refuse either
The
attitude of the
to
kill
or to eat them."
Fang subtnbes
Guinea amounts almost
to a taboo;
of
In
1891,
it
Equatorial
they believe that
eating apes, especially gorillas, causes
women.'"
d'lvoire,
was reported
infertility in
that the Fang
woman seeing a gorilla

one.'^ Some Fang tribes hold
believed that a pregnant

risked giving birth to
apes
killing of
is killed,
to the
among many
The former peoples

head hunting
taboo
is
ones as
may
(the Iban
of
source
be used
still
'^'
Borneo who engaged
in
and other Dayaksl some-
much as human
energy
of spiritual
for their long-
house communities. The Batak people

believe that eating orangutan
them
eat
to
populations
Islamic
times valued orangutan skulls as
still
improve their
to
it
medicinal purposes and witchcraft,'"
of
meat
Sumatra
will
make
strong.'^'
Many peoples have valued apes as simply

another source
of
meat, which
is
"somewhat sweet,
but with a nice taste."'" This attitude has been
spreading
in
where
the cities of Africa,
systems have
belief
lost their
traditional
power, but where the
The
logging companies recruit their laborers.
taboos and the movement

people
of
who no
into the great
toss of
ape areas
longer respect these traditions
represents a significant threat to the great apes.
The
local
people near
bos for religious reasons

for sale at a tourist gift
babies
of
northern Nigeria], ape body parts

for
when
generally taboo,
is
to
areas
Sierra Leone, Senegal, Liberia. Mali, and
e.g.
(in
in
the powdered bones and hair are
bathwater
primate meat
thought
is
believed
is
Wamba
in
DRC,
for
example, traditionally refrained from hunting bono-
the gorilla to be their totem.
A skull
as
killed
Congo, the
In
DRC, even
In
strength and health. Although
spirits of their
60 ethnic groups have chimpanzees as
their
added
become human
trying to
is
Liberia, the
In
Basin, bonobos represent
and the meat
strengthen young children.^'
is
living
as not.
Guinea, chimpanzee blood
cure epilepsy,
to
in
her
apes being
of
belief
part of a circumcision ritual for young
'"'
men.^
descendants; and
his or
and
Kwele, Kota. Mboko, and Djem peoples eat gorilla
meat as
the Batang Ai area of Sarawak, the orangutan
form can help
many cases
just as
a result of tradition
shop
in Bali.
'"
but, in the
mid-1980s,
poachers were recorded hunting bonobos

meat.'" The 'bushmeat
Africa
IS
crisis' in
for
West and Central
new
a reflection of the spread along the
logging roads of a cultural and market system that

enthusiastically
IS
consumes smoked ape meat;
this
penetrating deep into areas where people once
held very different beliefs. Nevertheless,

ple
these regions
in
still
many peo-
believe that these hominids
can learn our languages, break sticks
Ithat
indicate their direction
forest
to
nests,
use
tools,
of
travel,
in
the
build
and develop their own cultures)
deserve better from the hominids with guns than

be
killed, gutted,
and smoked over a
to
fire,
Julian Caidecott
227
World Atlas
Box 13.4 BUSHMEAT HUNTING AND TRADE

SENDJE, EQUATORIAL GUINEA"
IN
been transformed
have
camps.
hunters'
into
Trapping on a subsistence level has been going
on around the village for decades, but the recent
The
Sendje
village of
continental Equatorial Guinea, hi
mam
of Its
in
Rio Muni, or
increase
km
by road south
activity (particularly in
situated
is
town. Bata, and about 10 l<m west of
the Ivlonte Alen National Parl<. During the country's
many
time as a Spanish colony,

area were employed
plantations, living
throughout the
Spam
logging or on
in
camps and
in
people
1968, these forest inhabitants
in
oil
palm
villages scattered
After independence
forest.
the
in
from
were
village of
Sendje
officially
came
into being.
Sendje's population has increased to around
400 people, but following the cessation
of large-
commercial logging and agriculture, job
scale
opportunities are
However, the recent
scarce.
discovery of offshore
oil
Equatorial Guinea's
in
waters has meant that the country as a whole
has
tal<en
fuelled
an economic upturn. This
urban demand
bushmeat, creating conditions
including
commercial trade
lucrative
animals.^'
boom has
oil
meat and
for fresh
In
addition,
meat
the
fish,
for
in
many
of
it
is
of
the
the fields
in
cassava and peanut
hunt and trap
income. The majority
the forest for
in
meat caught
sold
is
in
the markets of Bata.
Sendje
is
of forest that
is
it
still
is
suited
to
com-
the gateway to a large area
relatively rich in wildlife. Old
logging tracks lead into the forest, enabling easy
access;
abandoned
The quantity
both
of
villages
and logging camps
bushmeat harvested
to its availability
and the cost
sources (see also Box
13.4).
example, has contributed

in
decline
Conversely, the
of
in
228
West
related
by
Africa, for
the decline of fish
nature reserves
mammal
more
in
in
Ghana, and
populations there."*
fish that is available in local
markets, the less bushmeat
The trade
is
other protein
turn has contributed to increased
hunting pressures
to
to
of
Factory fishing
foreign vessels along the coast of
stocks." This
hunting
now
of
takes place well inside Monte Alen
means
National Park, which
a trek of up to 30 km,
or 10 hours, from Sendje.
Trapping
much more common
is
than hunting
with guns, as the entry costs are lower (wire versus

a
shotgun and expensive, unreliable cartridges)
and fewer
skills are
needed. Around 75-80 trappers
and hunters were recorded during

study With a single exception
their
ages ranged from 13
all
in
15
month
were male, and
Ih years. Only around
to
15 of these people used guns, and this
was
usually
conjunction with traps. More than 5 000 active
traps were recorded
with
mean
younger,
more
the area at any one time,
in
100
about
of
per trapper;
some
and more commercial
vigorous,
or because they enjoyed

only
means
is
Mammal
earning
involve about 3
it,
but
because
it
was
their
money
surveys
and medium-sized
show
that prey
has been
species such as
terrestrial
duikers (forest antelopes! and large rodents that

are
targeted
by
trapping,
low throughout
are
Primate species are also targeted by
hunters with guns; primate density
is
much more
dependent on the distance from the

therefore
the
intensity
Primate numbers are

farthest
camp
hunting
million tons of wild
'West Africa, with
of
still
village
hunting with
quite
(inside
Monte Alen
in
Central and
levels in rural areas
ranging up to about 16 kg per person per
and
guns.
healthy at the
the
meat annually
consumption
year.^'
similar consumption rate of around 12 kg per
person per year was recorded
mid-1980s," and
is
in
Sarawak
in
the
generally consistent with data
from other tropical locations where people eat
bushmeat. However, consumption rates
of
up
to
06
kg per person per year have been reported for
hunter-gatherers
sold.
bushmeat may
of
severely depleted near Sendje. Densities of small-
the area.
particularly well
mercial hunting, as
people
search
into the forest in
prey The majority of commercial trapping and
other rural villages
women work
main crops
men
trappers
to
they did not choose to trap or hunt through tradition
the country, while
cultivating the
deeper
of
led
trappers operated up to 250 traps. Most said that
locations. In Sendje, as
for subsistence,
has
of wild
in
meat from increasingly remote
profitable to trade
number
of
traps)
of
and trapping
hunting
of
terms
improved infrastructure and
reduced transportation costs have meant that
in
number
total
traveling ever
re-
located forcibly to settlements along major roads,
and the
and
scale
the
in
north Congo.'
In
in
the central
the Peruvian
Congo Basin and
in
Amazon, bushmeat
National ParkI, but abundance
is
much
lower near
the village. Most conspicuously, the black colobus
preferred over the cheaper and
uing
demand
now
is
absent around
virtually
total of 8
in
396 animals were harvested
2003,
in
which the majority were
of
ungulates 135 percent! and rodents (33 percent],

belonging
fresh
provision of alternatives
particular to two species: the blue
in
consumer markets, demand
is liable
only to increase.
and great apes,
made up
28 percent!. Primates
only 10 percent of
the animals taken, reflecting the relatively low and

still
recent use of guns. Only one chimpanzee and
one
gorilla
were known
have been
to
killed in 2003;
over half of the primates taken were black colobus.
areas where hunting
records, suggesting that the

still
is
number
to
travel
in
yet,
for
there
is
bushmeat
no active
hunting regulations Iblack colobus

for
example, are supposed
to
be
protected under Equatorial Guinean law! and huntprotected areas continues unchecked.
ing within
Unless urgent action
economic nor the
bushmeat trade
in
is
taken, neither the socio-
biological sustainability of the
Sendje appears
to
be achievable.
Noelle Kumpel
market
relatively rich in primates.
combined with the

need
taken or
Monte Men National
hunted animals recorded
of
of
As
intensive, black colobus
is
among animals
rarely feature
and without the edu-
cation of
the brush-tailed porcupine lAtherurus africanus.
large
the form of improved
in
portunities for rural people!,
enforcement
Park area
will
Without greater
fish.
the development of other income-generating op-
duiker [Cephalophus monticola, 30 percent! and
In
meat and
sustainable fisheries and livestock husbandry lor
the village.
Sendje
avail-
rise with contin-
economic development, more people
[Coiobus satanas], a large-bodied, easily hunted

species rated as Vulnerable by lUCN's Primate
Specialist Group,
more widely
As incomes
able frozen produce.
Smoked monkey,
Equatorial Guinea.
The
Noelle
Kumpel
overall,
hunters and trappers
fact that
ever farther from the village
in
pursuit of prey, indicate that hunting and trapping

at
such
levels
At
cannot be sustained.
present,
and chimpanzees are
gorillas
hunted only opportunistically
in
this
Both
area.
species are taboo for the Fang ethnic groups that
Sendje and. although Fang people from other
live in
parts of Rio Muni will eat
has a low value
ticular
gorilla
it,
meat
markets.
in local
In
in
par-
view of
the fact that encounters with a gorilla group can be
dangerous
for
humans,
injury
from
trap
is
currently a greater risk than the threat of being
shot by a hunter for gorillas

In
Bata,
bushmeat
is
over other types of meat or
has been reported
to
in
the area.
not necessarily preferred

fish,
but fresh produce
be consumed
at a
is
rate of
19-168 kg per person peryear.^^
and
The national and international trade

great apes also appears to have increased
years across
many range
are the availability of

the
live
bushmeat trade
Iprlncipally affecting
because
in
in
The main causes
orphans as a by-product
West and Central
of
Africa
chimpanzees and bonobos
hunting related
[affecting
to
orangutans!,
logging
in
Southeast
and poor economic
in
many countries"'"'
(see Chapters 16
17!.
Great apes are also frequently injured or killed
live
recent
gorilla infants usually die before reaching a
likely buyer!,
Asia
in
states.
conditions
where snares are
set for other species,
medium-sized ungulates."
projects
to
specific
particularly
in
programs
East Africa
Kibale National Park, Uganda'^' and
Virungas!."The presence
of
usually
Primate conservation
sometimes include
patrol for snares,
(e.g. in
^^
in
rangers dedicated
the
to
the removal of snares also deters other forms of

illegal exploitation of the forest.
229
World Atlas
ECONOMIC CONSTRAINTS
governments
Tfie
23 great ape range states
of tfie
generally been
tiave
supportive of conservation
efforts for tfie great apes, even ttiougti ttieir re-
sources are very limited, especially

Indonesia. This
sfiown
is
Africa
in
and
dramatically by the
ranking of these nations according to the
Human
Development Index (HDII
Nations
of
the United
Development Programme (UNDP),'"

cepted indicator of
in
a widely ac-
and social wellbeing
vi/ealth
national societies (Table 13.5).
A low HDI rank
implies a challenging development context that

IS
affected by conflict,
and demand
poverty,
for
the extraction of natural resources, often at the
expense
of the
environment.
The HDI can be computed

and
A family
living
near
Kanyanchu on the
of the
for 177 countries,
23 great ape range states, 21 are ranked
between 109th (Equatorial Guinea) and 177th

(Sierra Leone). The only exceptions are Liberia,
forest
boundary, Kibale
which
59th.
Nicole
is
not ranked, and Malaysia, which
The
last figure
Simmons
may
is
ranked
mislead, as Ivlalaysian
Borneo ISabah and Sarawak), where orangutans

live,
Table 13.5
Human Development
Index
rankings, 2002"'
V(/orld
rank'
is
Sabah and Sarawak are constitutionally responsible for managing their own forests and wild-
Ape range
state'
life
without necessarily receiving federal support to
59
Malaysia
do
so.
109
Equatorial Guinea
the range states have significant challenges that
111
Indonesia
can
122
Gabon
131
Ghana
139
Sudan
The HDI analysis indicates
make
it
difficult to
that virtually all
undertake the organized,
long-term social investments demanded by successful conservation,
least
at
according to the
prevailing model.
141
Cameroon
U4
U6
Congo
range states, and most protected areas include
Uganda
some
151
Nigeria
these protected areas are of world-class quality,
157
Senegal
159
Rwanda
160
Guinea
Great apes are protected by law
having
in
great ape habitat and populations.
all their
Many
of
been selected as World Heritage Sites
and/or Biosphere Reserves. They
permanent
setting aside
in
all
represent the
law of large areas that
162
163
Cote
166
Angola
168
Democratic Republic
169
172
Guinea-Bissau
implies additional economic hardship. To judge
173
Burundi
from events, the commitment
174
Mall
their protected areas
177
Sierra Leone
much wealthier donor community.

A single example is enough to make this
particular point. A plan for the establishment and
a Out of
b
might otherwise have been used for logging,
d'lvoire
farming, mining, or other purposes, at significant

opportunity cost to the countries concerned. This
ot ttie
Congo
77 countries.
Insuffici ent
data to allow calculation for Liberia,
reveals the willingness
of
these
societies
to
preserve their national patrimony, even where this
development
230
and more rural than
significantly poorer
the urbanized Peninsular Malaysia. Despite this,
of
may
far
of
range states to
exceed that of the
Cross River National Park
in
south-
Box 13.5 EBOLA
AND GREAT APES
IN
IS
CENTRAL AFRICA
now
may have spread
feared that Ebola
The worst-case scenario

Diseases that can be transmitted from animals
to
tiumans under natural conditions IzoonosesI have

been
we have shared
existence for as long as
in
the Earth. Through the centuries, diseases such as
of
infection during
Syndrome
Respiratory
Acute
(SARSI and Ebola outbreaks have taken their
human
hemorrhagic fever virus
in
on
only one of at least 100
is
humans and
infectious agents that
toll
The Ebola
populations around the world
Ebola was identified
human
times
In
in six
in
976."'
populations
Since then,
at least a
Central Africa,
in
small area
in
996." Tens of thousands
human outbreaks on
samples or on
wildlife
region
the
in
the collection of
observations
in
the forest to
determine conclusively whether or how Ebola

affected the ape populations.
Prior to the 2002-2003 Ebola outbreaks

anticipation
in
of
in
an epidemic or
it
disease event, the Wildlife Conservation Society
worked together with the European Union program.
gorilla,
chimpanzee, and human populations were

hard.'^' In a
994 and
to
Ebola
dozen
different countries of Equatorial Africa.
recent epidemic
human
of
No work was undertaken
Ebola.
Congo, and
has affected
great
and chimpanzees may have died from
during the
great apes have
common.
in
and chimpanzee populations came close
disappearing during outbreaks
of gorillas
Severe
for Ebola
the Minkebe forest region
in
northeastern Gabon, where western lowland
gorilla
have crossed from animals to humans. Recently,
zoonoses have captured international attention
hit
northwestern Congo, this
and
Conservation
systems
management
areas
Forest
of
(ECOFAC),
staff of national
Congo and Gabon
in
Use
Rational
Central Africa
in
in
to
Eco-
tram
parks and protected

techniques for con-
devastating disease killed over 130
humans and
ducting wildlife censuses, monitoring the health
was estimated
population of
of wildlife,
about
to
have
killed half of a
200 great apes. The absence
family groups of gorillas
of
entire
and chimpanzees during
conducting postmortem examinations,
and performing standardized data collections.
The results demonstrated mortality
of
gorillas
and following the outbreak was confirmed by
due
laboratory testing of samples collected from gorilla
linkages; the similarity of the virus afflicting both
Some
members of
even
carcasses.
gorillas
other
the group had died of Ebola
hemorrhagic
fever,
chimpanzees exposed
east Nigeria
of
apes may have arisen
the bubonic plague, rabies, tetanus, and measles,
because
into the
Odzala-Koukoua National Park."
WWF-The
survived
as had
to
been observed
Ebola
was prepared
after
in
Cote dlvoire.^'
in
with the joint support
the government of the United Kingdom, and the
most
state,
of the
'"
comprises
Gorilla
gorilla
Development Credit Agency
(Kreditanstalt
Wiederaufbaul visited Cross
River state
fur
June 1991
in
to
assess the
it
divisions:
Oban
Okwangwo
Pan
(containing
diehli].
The plan was
accepted by the Nigerian federal government
the
and the German
paration for putting
two
that there
continued overleaf
Cross River
in
for multiple,
same time." This suggested
amend
Nigeria-Cameroon chimpanzees and Cross River

gorillas.
forest at the
the project and to
(containing Nigeria-Cameroon chimpanzees,
troglodytes vetlerosus] and
humans; and evidence
great apes and
genetically distinct Ebola viruses circulating
" The park contains
remaining moist forests
and
Ebola hemorrhagic fever; direct genetic
It
Global Conservation Organization,
European Commission."
to
its
feasibility of
structure
in
pre-
out to international tender
Consulting firms were shortlisted for the manage-
ment contract
awarded
in
in
March
More than
beginning
993, and the contract
was
October 1993.
five
of project
years elapsed between the

planning and the beginning of
in
project implementation. While this story gradually
December 1989, which then asked the European

Commission for assistance to implement
In April
unfolded, the Nigerian government proceeded to
it.
1990, a draft financing proposal
the European
was prepared
Commission delegation
in
by
Lagos, and
was subsequently approved by the Commission in

Brussels, A team from the European Commission
fulfill its
stated intention of creating the Cross River
National Park.
political
It
attempted coups
strikes,
achieved this despite
and economic
difficulties,
d'etat,
factional
its
own
which included
riots,
general
and financial crises on an enormous scale.
231
World Atlas
might be multiple reservoir species. These trained
teams drew on
field
essential, established
rela-
in
humans,
Africa,
tionships with local villagers and hunters, to enable
link
them
gorillas or
to detect
and report Ebola
apes months before the
fatalities in great
human
first
cases. Since
the mid-1 990s, the earliest cases of Ebola infection
Ebola outbreaks
in
prior to wider outbreaks
in
Central
have been analyzed. These have shown a
between
handling
the
chimpanzees and
Ebola-infected
of
susceptibility to the
disease. Reducing the frequency of this contact
route
reduce
could
the
incidence
of
human
Central Africa
12E
Species
Central chimpanzee
Western lowland
gorilla
Ebola outbreaks affecting apes

Year 1994-1997
Year 1996
ED
Year 2001 -2002

Year 2002-2003
RiOMUNI
'
EQUATORUL
|
GUINEA
GABON
50
^-^
12'E
precisely on
Can great apes be saved from Ebola^ Sc/ence 300:
of international activity,
it
approved Cross River National Park
Rather than investing solely
in
October
legally created the national park
on October
2 1991.
were
less acute than
assessed them
by the park
was
to
the project area.
planning for early
implementation, a program might instead have
been undertaken involving basic conservation work

priorities,
which were and
remain: "to protect the forest; to maintain lines of
WWF
communication between the people affected by

that protection and the people doing the protecting;
undermine
local trust
in
the conservation process as financed by outsiders.
WWF
in
in
and the most significant
had been thought when
initially,
effect of the delay
Had
expectations
around the obvious
Fortunately, the threats faced
16^5.
would have been possible
to avoid raising
1989, leading to Presidential Decree No. 36 of 1991
232
G. (2003!
schedule, the Federal Nigerian Council of Ministers
which
1501(111
14"E
Based on Vogel.
Despite these interruptions, and
100
,'
been better prepared
for the
slow pace
and
to help both sides
limits of their
understand the ecological
environment and how
to live
the best
possible lives without exceeding those limits.""
outbreaks, while also discouraging the hunting of
Work
underway
IS still
to identify the natural
Some
resen/oir lor reservoirs) of the Ebola virus.
species of
fruit
and then shed the virus
excrement.'" Fieldwork
in
wildlife
need
of
following
the
apesi
great
be addressed:
to
anticipate Ebola outbreaks and populations at
their
in
CAR has demonstrated
number
some
to
provide better support to areas
teams
establish monitoring
rodent species."" Similar
of
Congo has found the same
order
risk in
that could be affected by the virus;
the presence of at least fragments of Ebola viral
to
determine the
bats.'^
existence and progression of the Ebola virus
These findings should be interpreted with caution as
and other serious infectious disease agents
the techniques used determine only the presence of
in
the genetic material; this does not always indicate
[affected species, mortality rates, resistance,
work
in
the presence of
live
apparently healthy
of
Ebola
widespread
IS
persists
It
at
it
people to reports of the presence
improve knowledge
response.'^""
ween and
and
evaluate
in
more
Ebola virus to other,
at forest peripheries, in
may
fragments, and
species or the type of habitat
which transmission between species
may
of
reducing the effect of Ebola
in
is
more
of
affected species
time and space, develop-
of
in
approaches
populations
li.e.,
to
strategies
for
preventive
of
disease
risk,
by
medicine
and
improve local community
human
view to reducing
hunting of great apes.
diseases.
contact with and the
order to understand better the disease
In
caused by the Ebola virus and
to
civil
of great
had
ape range states since 1990, and
Map
Leone, Liberia, and Cote
on
impact
13.11. In
have occurred
many
for
have greatly affected a
conflict
significant
populations (see
William Karesh and Patricia
develop methods
AND CONFLICT
POLITICS
Wars and
conflicts
meta-
education and awareness campaigns, with a
populations and promote the transfer of the virus,
have
of
connected
using Ebola hemorrhagic fever as an example
likely
as has been observed with other
number
are
improved hygiene practices; and
in
between animal
viral
management
that
dispersal across the landscape), and other
in
climate or vegetation patterns
alter ecological relationships
reservoir species and
grams, separation
ment
be more
mosaics."" This could reflect the preferred habitats
occur Changes
within species, strains, immunity!;
ways
using techniques such as vaccination pro-
The conditions
level.
a virus of deep forest refugia, Ebola
to
its
transmission bet-
probably causes minimal disease
the population
of the reservoir
Ebola virus and
of
and other infectious diseases on great apes
vulnerable, species are unknown. Rather than being
common
of the
mode
ecology Ireservoir,
date suggests that
Equatorial Africa
Ebola
of
hemorrhagic fever and other diseases;
more natural
the transfer of
for
to
believed to have one or
is
which
in
problems
in
establish response plans to alert appropriate
Central
nature between observed outbreaks
in
humans.
hosts,
evidence
in
the forest, and their impact on wildlife
natural barriers, etc. I;
in
exposure having been
immune
successfully overcome by an
scientific
antibodies to Ebola
humans and primates
Africa, indicative of previous
The
in
or viable virus. Other studies
have shown the presence
in
on
other diseases
objectives
humans and
in
understand and prevent the effects
bats and insectivorous bats can sur-
vive the infection
particles in a
spread both
to prevent its
las well as to
and chinnpanzees-
gorillas
in
West
local
Africa,
ape
major
Guinea-Bissau, Sierra
d'lvoire; in
DRC, Congo, Angola, and CAR;
in
Central Africa,
East Africa,
in
years.
availability
homes
and
production;
levels
of
and
survive.^''
conflicts
increase the
guns, displace people from their
farms,
all of
and
reduce
agricultural
these factors can increase hunting
illegal
"
Armed
Reed
Ivluch
logging as people struggle to

of the
money
for international
comes from bilateral grants, funds

may be frozen when security deteriorates. This
conservation
that
Sudan, northern Uganda, Burundi, and Rwanda;
can close conservation projects and cause the
and
loss of experienced project staff (e.g. as
in
Southeast Asia,
in
Indonesia, particularly
in
the Virungas
northern Sumatra where Acehnese separatists
in
have been battling the Indonesian government
often been
in
happened
the mid-1990s). Attempts have
made, however,
to
maintain at least
233
World Atlas
Low-intensity conflicts are those of fewer ttian 25 deattis per year;

intensity conflicts are ttiose of
conflicts at the
Map
same
in
1 000 deaths per year IVtultiple

symbols overlaying each other.
13.1 Historical conflicts affecting the African great
some
activities at great
ape sites during periods
of
conflict."''"'"
War
armed
toll
kills
conflict
in
conflict,
animals
Rwanda, three
for
refugee
of the four
in
camps
and destruction
the Karisoke Research Center
in
of the
Conflict diverts revenues to

in
in
DRC
workers
at
gorillas."
"
arms and away
conservation or poverty
relief,
and deters responsible private investment.
Some
less scrupulous
companies take advantage
of the
chaos
valuable resources,
to extract
as a
some-
to
means
political
engage-
encourage the extractive industries

liquidating
of
financial gain.^'This
is
natural resources for
disastrous for great apes and
their habitats, as well as for people (especially the

poorl, although the stifling
of
economic
activity,
depopulation, and chaos brought by conflict can

occasionally relieve short-term pressures on the
environment
by,
example, deterring logging
for
companies.
The long-term impacts on great apes
park infra-
Volcanoes National
many mountain
from social investment
in
or near to the
structure, and to the deaths of several
Park, as well as of
wood
bushmeat. During the
Virunga National Park. Subsequent conflict
23^
conflict.""' (Military
ment impoverishes nations and may tempt gov-
North Kivu, DRC, were located
led to looting
maintaining a
ernments
unrest have tal<en their
political
gorilla habitat by uncontrolled harvesting of

of
in
great apes as well as people, and
can often put massive pressure on forests and
and
times participating with local military and
and
Refugees, displaced as a result of
for fuel
ape range states
actors
on both the eastern lowland and mountain
gorilla.
war
fiigti-
more than
location result
recent wars
in
of peaceful conditions,
increasing trade
more
traffic in
in
however,
profit,
is likely to
lead to
most areas, which may allow
bushmeat. The purpose
would then be expected

to
of the
Central Africa are unclear The return
to shift
of
hunting
from subsistence
but would keep ape populations under
pressure. Peace
an expansion
of
may
bring other dangers, including
industrial-scale logging, mining.
promoting
and forest clearance
for farming,
all
access and the spread
of fiunting to
new
Less violent
areas.
change can also impact
political
ape conservation adversely. For several years after

the
fall of
servation
the Suharto government
deforestation increased markedly, even

areas."'
"
1998, con-
in
Indonesia virtually collapsed, and
in
''"'
Governments
commitment, or capacity
political will,
effective conservation
work, which
as a high priority This
is
is
to
sufficient
undertake
seldom seen
especially true
torn states. Sudan, for instance,
emerge from
protected
in
rarely have
is
in
war-
beginning to
a 30 year conflict that has killed or
displaced millions of people, and the country
expected
be engaged
to
is
national reconciliation
in
and rebuilding for the next few years, although the
and ethnic cleansing
conflict
province of Darfur that erupted
in
In
western
the
200^ may disrupt
these plans. Conservation of that country's remnant
is
not likely to be viewed as
and Sudan has
a priority in these circumstances,
not yet opted to join the Great
many
Politicians in
beginning
sources
Project.
developing countries are
address the impact on natural re-
to
of
Apes Survival
mushrooming human populations. The
great ape range states contain countries with

of the largest (e.g. Nigeria, Indonesia] or
(e.g.
some
densest
Rwanda! populations on Earth, and some with
the shortest
expectancies
life
(e.g.
Cote
the lowest gross domestic product
Poverty
is
rife
many
in
of
d'lvoire)
and
Burundi).
le.g.
these countries, which
places additional burdens on governments and the
management
of natural resources. Ultimately, the
apes
fate of the great
people
in
their
is
linked to the future of the
range states, and particularly their
capacity to stabilize their populations and

their social
William Karesh
one
of
the
few success stories
great
of
conservation - the use of great apes
in
ape
tourism.^^
Close contact with a group of habituated gorillas

or orangutans
will
pay
of
considerable
to
samples
amounts;
tourists
for
Ebola diagnostics
an experience for which tourists
is
however, travelers and are therefore most
expose apes
A conservation team
collecting
in
Congo.
are,
likely to
new pathogens. A number of cases

known among the mountain
cross infection are
gorillas of the Virungas, including an outbreak
1988 suspected
lar
to
morbillivirus,
have been
which
of
measles or
killed
six
habituated
females;'" an epidemic of bronchopneumonia

Elizabeth
in
a simi-
in
A Williamson
Goldminers
in
Nyungwe
National Park, Rwanda.
meet
and economic development needs.
These solutions
will
need
to
be based on wise use
natural resources and respect for other species
of
if
great ape populations are to survive.
DISEASE
As closely related animals, humans and great
apes can
infect
one another with a wide range
of
diseases and parasites. Being far more mobile and

in
contact with a global spectrum of pathogens,
humans
are
much more
likely to
introduce illness
to small, isolated
populations of apes than the other
way round. The
potential for transmission
opposite direction
is
however
a strong
in
the
argument
against the consumption of primate meat.
Disease transmission also risks undermining
235
World Atlas
1990, which affected 26 of the 35 gorillas
had been exposed
that
gorillas;" and
debilitating
members
affected all four
tourists,
to
sl<in
two
disease that
group which
of a gorilla
had been habituated for tourism, and
them.
a group
in
l<illing
l<illed
one
of
was
loss
minerals and
of
and degradation
oil
causes habitat
apes are
a limited area;
in
mainly affected by the associated infrastructure
Roads and pipeline routes allow
development.
access previously
hunters
to
contracted from exposure to local people rather
Oilfield
workers and miners frequently consume
than to tourists."'
bushmeat, and may
In this last
case, however, the infection
MINING AND OIL

The extraction
The continued
numbers
of
daily
exposure
people and their diseases
major potential threat
apes
of
to
is
to large
considered
great ape
tourism
sell
on
it
incomes. Major mines and
exploited forests.
little
supplement
to
have the virtue that they tend
least
by large companies which,
if
their
developments
oilfield
to
be
they wanted
have
to,
operations,'^ although this has not yet jeopardized
the capacity to regulate hunting and wildlife trade
the sun/ival of whole populations of great apes.
their areas of operation.
In
in
hemorrhagic
contrast, Ebola
most
virulent viral diseases
50-90 percent
of
all
fever,
one
of
the
known, causes death
afflicted
humans and
The coltan boom
damage
to
black alluvial ore
among
which
gorillas." This virus
976 after outbreal<s
Sudan.'"
the
It
is
mammals. Ebola
linl<ed to direct
was
northern
first
identified in
DRC and
southern
transmitted by direct contact with
body
blood,
in
fluids,
and tissues
of
infected
infections in people have
been
contact with gorillas, chimpanzees,
monkeys, forest antelopes, and porcupines found

dead
in
map
is
based on GLOBIO
medium-high
loss,
and yellow
2
a
found
in
of
'" '"
Coltan
loss.
is
columbium and tantalum,
riverine deposits
Central Africa.
in
After the tantalum has been extracted,
it
is
used
electronic capacitors, particularly those found
in
in
miniaturized equipment such as mobile telephones,

laptop computers,
panning,
like
and games consoles. Coltan
gold panning,
is
often undertaken by
individual freelance artisanal miners.
in tropical
in
DRC
is
Africa
analyses for the year 2000 Black indicates a
low-medium
provides an example
Between 1998 and 2003, the war
the rain forest Isee Box 13.51.
Figure 13.1 Impacts of infrastructure development

This
is
DRC
in
process that caused great
great ape populations.
apparently produces an even higher mortality rate
in
of a poorly controlled
at
managed
likely
high loss
Green areas experience low impact
in
species diversity, red a
of infrastructure
development.
^^î^pSP^?!^
k.
:3
.--"
'
m^^B
^"! ^^HjUw.?
^m
^H
^H
High
human Impact
Medium-high human impact
Low-medium human impact
236
is
...
."..^
m
m
"<
1Im
1
w
F
fl
i
Ik
1
J
4
Sfft?W-l^
believed
have
to
killed
some
the effects of
displacement. This coincided
a global
in
the coltan market
2001.
in
In
in
vtfith
2000, followed by a
DRC, the boom
Electronics companies
people,
million
either directly or indirectly from
boom
slump
led to a rush of
popular consumer products

wildlife
in
There are
Kahuzi-Biega National Park was
mineral.
slaughter
to
Reserve by some 3 000. These included inde-
now moves
and trade
fessional hunters
armed with automatic
Kahuzi-Biega National Park,
in
feeding the miners on local wildlife.
most
that
of
in
rifles
many
insecurity
to
concluding that the war
to
itself
was
the armies of neighboring countries
ation of
in
fuelled
and control the production
DRC under
a mining code
in
to
fewer than 4 000 animals, but
the region has not yet allowed surveys
confirm this decline.
Great apes are often confronted simultaneously

by multiple threats.
by
were noted
effect
resources.'^''
'^^
Assessing their combined
can be challenging, as these threats often
interact with
to
or benefited from the exploit-
DRC's natural
difficult
Multiplying threats
global attention,
revenue from coltan mining. Various companies and
have participated
is
THREATS AND POSSIBLE FUTURES
series of reports from the United Nations
brought the situation
it
ends up. There are
other animals and
Security Council and nongovernmental organizations
to legitimize
of coltan in
around 17 000
suspected
the park were eaten, along with
the elephants, and
birds.
is
this material
developed by the
the then population of 8 000 eastern
lowland gorillas
all
It
Congolese coltan."
DRC government and the World

Bank. During the coltan boom years, the eastern
lowland gorilla population may have collapsed from
By December 2000, 300 pro-
were working
both the war and the
not well audited so
is
assess where
pendent workers, their families, and others working

labor
to
DRC. Many have since declared
exports from DRC, however, and the
still
supply chain
occupied by about 10 000 people, and Okapi Wildlife
as forced
in
that they will avoid purchasing
miners
those poorly protected national parks that hold
the
were taken unawares
by public dismay at the reported contribution of
each other as well as with the apes
themselves. The simplest way to assess the level
of
threat posed by a combination of pressures
to
is
Figure 13.2 Infrastructure development projected for the year 2030

This image reflects
GLOBIO
2 scenarios.
Black indicates a
and yellow a low-medium loss Green areas are
High
likely to
likely
high loss
in
species diversity, red a medium-high loss
experience low impact
of infrastructure
development.
human impact
Medium-high human impact
237
World Atlas
Figure 13.3 Impacts of infrastructure development

This
map
is
based on GLOBIO
medium-high
loss,
and yellow
low-medium
add togetfier the pressures
in
at
can be done by overlaying
each
digital
various pressures over the entire
number
to
combined score
be calculated.
to
fragment habitat areas, and

transport of
of
range
the
of
One study employed
a ranking
system from
compare the
current situation for various pressures
in
three
development.
facilitate
both
the
bushmeat and access by poachers and
settlers.""
""''
Outside the few reserves with effective law
enforcement, the only places where apes are

relatively safe are in very
where there
or places
lleast threat! to k (greatest threat) to
species diversity, red a
in
of infrastructure
maps
serious pres-
of
high loss
location. This
be assessed or ranked, and a
sures at a location
likely
Green areas experience low impact
loss.
species, using geographic infornnation system IGISI
software. This allows the
Southeast Asia
analyses for the year 2000. Black indicates a
is
remote areas, swamps,
powerful local support for
protection, such as in the Lac Tele/Likouala-aux-
Herbes Community Reserve
in
northern Congo.
'^'
Habitat loss mainly takes place through agricultural
and
parts of CAR. Logging, mining, hunting, agriculture,
expansion
and human presence were calibrated and the
logging for timber and pulp, and around mining
results
were summed. The combined threat
for the
two sectors
Park, Dzanga and Ndoki, and the
area were determined to be

respectively.'^
moderate
threat
in
levels of logging
the
Dzangha Sangha
1.8,
1.5,
and
Roads were considered one
most important threats
levels
Dzanga-Ndoki National
of the
in
all
l.U
of the
three areas, with
and mining also posing a
Dzangha Sangha
burning
These
operations.
along
together
roads,
extractive industries, result in
road
corridors,
with
the
'boom towns' without
sufficient local food supply; this leads to increased
demand
ficant
for
bushmeat, which represents a
signi-
income-generating opportunity for families
with the lowest incomes."'
between roads, extractive
^"'
"'
The relationship
industries,
and increased
bushmeat trade has been confirmed by numerous

'"' "''"""'"
area.
studies."'
The mapping
Infrastructure development
Roads play a central

of tropical
role in the loss of great
moist forest biodiversity
in
apes and
general. They
provide access to mining and logging companies.
238
of
road networks, settlements,
and mining operations can help
to indicate
areas of
probable habitat loss and the degree of exposure

of great
apes
to hunting.
On
this
assumption, the
t'^j
High
human impact
human impact
IVledium-high
Figure 13.^ Infrastructure development projected for the year 2030

This image reflects
and yellow
GLOBIO
low-nnedium
2 scenarios.
loss.
Black iniiicates a
Green areas are
GLOBIO computer model was developed

United Nations Environment
map
help assess and
human
area
land
abundance
of
living
is
used
reduced
with
to
to
estimate the
biodiversity
organisms following
structure development.
and
infra-
can also be used
It
predict the impacts of proposed
to
developments by
helping to visualize the zones around roads, major

trails,
to
settlements, dams,
be a reduction
Four zones
the
in
etc.,
where there
abundance
is likely
of wildlife.
'"
impact on biodiversity are defined
of
likely
high loss
using this tool:
in
species diversity, red a nnediunn-high
The high-impact zone denotes
a belt
generally heavily used by people, and
is
the area within which
than 50 percent of
by
more than
all
more
recorded species decline
few great apes. Medium1-3
km
medium-high impact
(i.e.
the area within
of all
recorded species
high-impact zones
to
from roads and settlements. This
is
and
is
where logging
is
known
to
usually most intensive; great apes are
decline dramatically
in
areas subjected to intense
logging, largely as a result of
hunting."""
chimpanzees, and orangutans have
all
these relatively intact forests are often subject
recorded species decline by
low impact (areas
population densities."
beyond the above
^'
'' '"
intensively
km
It
of
is
roads
Due
meat through the
hunters will move beyond 10
3-8
and
falling
to
to
and logging generally contain much higher ape
most
zones).
Low-
medium-impact zones occur 3-10 km from roads;
1-25 percent
percentl;
Gorillas,
been shown
use food resources within logged forests, pro-
carry loads of
more than 50
lie
a typical
operating radius for log-skidding [dragging logs to

their initial destination),
more than 50 percentl;

low-medium impact (i.e. the area within which
decline by
of all
is
charac-
heavy hunting pressure. Areas free from hunting
50 percent);
which 25-50 percent
wide
and very
terized by logging, farmland, settlements,
to
(i.e.
km
surrounding roads and towns; such an area
viding that they are not being hunted." "
high impact
loss,
experience low impact of infrastructure development.
for the
Programme (UNEP)
the environmental impact of
development.'^" This
of
likely to
in
km
hunted zone
is
to the
need
forest, very
to
few
from roads; the

typically within
logged areas.
possible to simulate future changes
distribution of these impact
in
the
zones by using simple
239
World Atlas
Red List
Table 13.6 Projected great ape population declines, 200^
such habitat
will
less than
percent remaining
be about
category by 2030 (see Figure
Timeframe
Taxon
[Gorilla
Pan troglodytes
be
IP.
(.
lost
THE ROAD AHEAD

Some great ape taxa
""
2000-2060
50 percent'
more threatened
are clearly
schweinfurttiii]
than others, but

ted
a
will
by 2010."
50 percent"
2000 + 3 generations
troglodytes]
Eastern chimpanzee
These figures are
orangutan habitat within protected areas
of the
benngei graueri]
Central chimpanzee
1
'
50 percent'
the low-impact
in
13.i|.
consistent with published estimates that A7 percent
Projected rate of decline
2000 + 3 generations
percent per year, with
Decline
is
projected based on recent major losses
have experienced or are expec-
all
experience sufficiently worrying
to
the Highlands sector of the Kahuzi-
in
merit
to
inclusion
as
Endangered or
declines
Critically
Biega National Park,

b Decline
is
projected based on habitat loss
in
East Africa and instability
in
DRC
Endangered on the
leading to
lUCN Red
200^*
The
List.^''
threats to great ape survival are difficult to tacl<le
continuing hunting pressure
isolation.
Although some excellent models
servation exist and there
assumptions about the growth and dispersion

infrastructure, for
in
example
that
human
areas with large
it
will
of
expand fastest
populations, close to
Less than 30 percent
to the following conclusions.'"

of the
is
led
habitat of each of the African great apes
currently classified as under tow impact from
the indirect effects of infrastructure development

(Figure 13.11.
of
The future annual
such habitat was projected
rate of degradation
to
exceed 2 percent
per year, with 10 percent or less of their habitat
remaining
the low-impact category by 2030
in
(Figure 13.21.
These results suggest
habitats will decline
rapidly
in
that great
ape
coming years
if
tal<e a
huge
ment
them.
is
infrastructure
development (see Figure
13.3).
Future scenarios suggest that the annual loss of
of
will
more
of the
many
Good manage-
countries.
Some argue
that protected areas are the best
targets for conservation investment as they are well
defined units that already exist

great apes
in
national law.
and a more inclusive strategy
is
needed
to
ensure
their survival in the broader rural landscape.
term solutions
the
Ivlost
outside protected areas, however,
live
rely
on a change
in
attitudes
populations and governments
range states. This
great ape
of
shift in thinking is
Long-
among
required to
hunting and land-use change within great
limit
of
it
crucial for the ape populations living within
ape habitats, and
as subject to low impact from the indirect effects
body
Protected areas form the core of the conservation strategies of
percent of orangutan habitat
currently classified
ensure the survival
effort to
in
con-
endangered ape species and subspecies.
current trends continue. Meanwhile, less than 36

is
a substantial
research on the pressures and their impacts,
natural resource concentrations, near coasts, and
around existing infrastructure. This analysis has
is
of
to
ensure that conservation and
development resources are available and appropriately
co-managed. Conservation
is
the
theme
of
the remainder of this volume.
FURTHER READING
Brashares,
J.S.,
Arcese,
P.,
Sam,
declines and fish supply
in
M.K., Coppolillo, P.B., Sinclair, A.R.E., Balmford, A. (20041
West
Africa.
Formenty, R, Boesch, C, Wyers, M., Steiner, C, Donati,
among
wild
chimpanzees
Bushmeat
hunting, wildlife
Sc/ence 306: 1180-1183.

F.,
living in a rain forest of
Dind,
Cote
F.,
Walker,
d'lvoire.
F.,
Guenno,
B.L. 1199V) Ebola virus outbreak
Journal of Infectious Diseases 179 ISuppl
II:
S120-S126.
Galdikas, B.M.F., Briggs, N.E., Sheeran, L.K., Shapiro, G.L., Goodall,
African Apes. Kluwer
Harcourt, A.H. 11996)
Is
J.,
eds 12001)
the gorilla a threatened species?
Howshould we judge?
Harcourt, A.H., Parks, S.A. (2003) Threatened primate taxa experience high
to
2^0
the
lUCN Red
All
Academic/Plenum Publishers, Boston, Dordrecht, London, Moscow, New
List criteria. Biological
Conservation 109
111:
P.
Biological Conservation75{2]: 165-176.
human
137-U9.
vol.
York.
densities: adding an index of threat
Hayes,
Burge, R. (20031 Coltan Mining
K.,
Can Commit
How
Democratic Republic of Congo:
in tfie
Reconstruction
the
to
DRC.
(he
of
Fauna
and
Flora
Tantalum-using Industries
Cambridge,
International.
UK,
http://www.gesi. org/docs/FFI%20Coltan%20report,pdf. Accessed March 18 200A.

Huijbregts, B.,
De Wachter,
P.,
Obiang, L.S.N. Akou, M.E. 120031 Ebola and the decline of gorilla Gorilla gorilla and
,
chimpanzee Pan troglodytes populations
lUCN
120041 2004
lUCN Red
http://www.redlist.org.
Kalpers.
J.,
Williamson,
List of
the Minkebe Forest, north-eastern Gabon. Oryx 37: ^37-443.
in
Threatened Species. lUCN-The World Conservation Union. Gland, Switzerland.
Accessed February 28 2005.

E.A.,
A.,
Nzamurambaho,
A., Lola, N., Mugiri, G. 120031 Gorillas in
the crossfire: population dynamics of the Virunga mountain gorillas over the past three decades. Oryx 37
131:
326-337.
Leroy, E.M., Rouquet,
African wildlife.
Formenty.
P.,
P.
(20041 Multiple Ebola virus transmission events and rapid decline of Central
Sdence 303: 387-390.
Nellemann, C, Newton, A. (20021 The Great Apes - The Road Ahead- UNEP/GRID Arendal, GRASP,
UNEP-WCMC.
http://viWW.unep-wcmc.org/resources/publications/GrASP/GRASP_5_complete.pdf. Accessed November 28

2004.
Peterson, D. (20031 Eating Apes. California Studies
Redmond,
(20011 Coltan
I.
Boom,
Gorilla Bust.
in
Food and Culture,
6.
University of California Press.
report for the Dian Fossey Gorilla Fund and Born Free Foundation.
http://wv;.bornfree. org.uk/coltan/coltan.pdf. Accessed
March 18 2004.
Robertson, J.M.Y., van Schaik, C.P. (2001) Causal factors underlying the dramatic decline
of the
Sumatran orangutan.
0^7^35: 26-38.
ThibauU, M., Blaney. S. (20031 The
Conservation BiologyM
UNDP
(20041
Walsh,
P.,
industry as an underlying factor
oil
Human Development
A.K., Kilbourn, A.M.,
Lahm,
R.,
De Wachter,
S.A., Latour, S., Maisels,
Starkey, M.R, Telfer, P, Thibault, M., Tutin, C.E.G.. White,
western equatorial
Wilkie, D.,
Shaw,
E.,
Africa.
Rotberg,
the
bushmeat
crisis in Central Africa.
Report 2004. http://hdrundp.org/reports/global/2004/. Accessed November 2004.
Abernethy, K.A., Bermejo, M., Beyers,
Toham,
in
1807-1813.
161:
P.,
F.,
Akou, M.E., Huijbregts,
Mbina, C, Mihindou,
L.J.T., Wilkie, D.S.
Y.,
B.,
Mambounga,
Obiang, S.N.,
D.I.,
Effa. E.N.,
(20031 Catastrophic ape decline
in
NatureUll: 611-614.
F.,
Morelli, G.. Auzel,
P.
(2000) Roads, development and conservation
in
the
Congo Basin.
Conservation Biology 14: 1614-1622.
MAP SOURCES
Map
13.1 Conflict data are
Buhaug,
H.,
based on the following sources:
Gates, S. (20021 The geography of
Gleditsch, N.R, Wallensteen,

dataset. Journal of
P.,
civil
war Journal
of Peace Research 39
Eriksson, M., Sollenberg, M., Strand, H. (20021
Peace Research 39
(51:
[41:
Armed
417-433.
conflict
1946-2000: a
new
615-637.
ACKNOWLEDGMENTS
Many thanks
Redmond
on the draft
WCMCI
to
Thomas
Butynski [Conservation Internationall, Alexander Harcourt [University of California. Davisl. Ian
[Ape Alliance/GRASPl, and David Woodruff [University of California, San Diegol for their valuable
of this
chapter Thanks also
to
Tonya Lander, Nigel Varty, Sarah Ferris, and Valerie Kapos
comments
(all of
UNEP-
for research into the literature supporting this chapter.
AUTHORS
Lera Miles.

Julian Caldecott,
Christian Nellemann,
Box
13.1
Box
UNEP/GRID Arendal
Alexander H. Harcourt, Department
3.2 Lilian Pintea.
of
Anthropology, University of California, Davis
Center for Primate Studies, Jane Goodall
Box 13.3 Julian Caldecott,
UNEP World
Institute, University of
Minnesota
Box 13.4 Noelle Kumpel, Zoological Society
of
London, and Imperial College London
Box 13.5 William Karesh and Patricia Reed, Field Veterinary Program and Wildlife Conservation Society
2A1
World Atlas
Chapter
Conservation
measures
play
in
Nigel Varp/, Sarah Ferriss, Bryan Carroll,
AND Julian Caldecott
emerging threats
The
forests
particular have
to
and
general
in
moist
tropical
apes
great
in
gone unopposed by
not
people and governments around the w/orld. A
measures have been discussed and
host of
implemented with increasing urgency and
v\/ith
apes
safe.
where they are
own lands as
share
in
to
do
forest refuges in the
hope
environmental benefits from the protected forests.
Meanwhile, the
community has
scientific
steadily
ments
kinds that aim to encourage
and their ecosystems, making
it
conservation planning, to regulate trade, and to
municate the unique attributes
of
promote transfrontier cooperation
ment
of
protected areas and
International
in
w/ildlife
nongovernmental
the
manage-
populations.
organizations
added
to
our knowledge
those
to
who
value them, and
decision
donors, and range-state governments have
chapter explores these varied
protected areas, improve their
engage with people

goal
in
is to
living in
encourage and enable improvements

conditions and
living
management, and
and around them. The
that protect great
the great apes
easier to
process telling
to
com-
these animals,
felt
by
achieve a
in
much
initiatives,
of the story of
in
human
sustainable relationship with the
from West
tropical moist forests of the Old World,
Africa to Borneo.
of
in-
PARTNERSHIPS
Widespread public
apes created
interest in great
a responsive
discussed
their deteriorating conservation status. Great
this chapter.
among concerned groups has
been tentatively extended

porations
the
in
include private corin
the timber, mining,
infrastructure sectors
pacted great apes

so
to
whose investments
energy, and
future.
in
the
efforts
fluence that support great ape conservation, as
Collaboration
to
great ape conservation. This
compliance with laws
apes and their habitats. These
stakeholder groups exert various kinds
in
all
guidance
providing
makers on how and where best
make investments
all
of
deepening the interest and excitement
(NGOsl, national NGOs, bilateral and multilateral
cooperated on interventions that aim to establish
of having a
revenues from tourism, as well as deriving
measures have included intergovernmental agreeof various
be
likely to
some communi-
so,
ape habitat areas have set aside their
ties in great
the investment of ever greater resources. These
have
all
im-
the past and are likely to do
Tourism ventures have been
environment
populations began to decline seriously
as
soon as projects involving
ape
the 1960s,
in
industrial-scale
logging, infrastructure, and plantation
development
entered their habitats. By the mid-1980s, the fate of

tropical rain forests
became symbolized
by a few
charismatic species, including the mountain
thanks
apes
media. The murder of Dian Fossey
to nature-oriented visitors for the benefit of
alarm over
for raising
established to promote the attractiveness of great
conservation. Sanctuaries for great apes confis-
to the influence of primatologists

in
gorilla,
and the
1985 high-
lighted at the international level the devastating
cated from hunters and traders have multiplied.
impact that poaching was having on the already
These are being used
small and threatened population
and,
242
to wild habitats
Where empowered
in
skillfully for public
education
places, to support the reintroduction of great
gorillas.
Populations of
all
of
mountain
the great apes were
under pressure from deforestation, farming, and
armed
hunting;
In
many
areas.
aggravated the situation
conflict
There was a growing sense
of
decade or more, during which time the fates
GRASP was
discussed
In
2001, and launched as a Type
lingness to pay for conservation action.
voluntary,
Starting
in
WWF-The
the early 1970s with the support
Global Conservation
Organization
(formerly the World Wildlife Fund, and

this In
North America) and the Wildlife Conservation
New
Society (WCS, formerly the

Society!
York Zoological
work
orangutan-rehabilitation
for
Bohorok and
in
called
still
research
field
the Leuser Ecosystem
NGOs soon
international
In
Ketambe
at
at
(both
Sumatra), numerous
turned their attention
to
play
of
several great ape taxa are likely to be determined.
worldwide public urgency, and an Increasing wil-
of
in
servation
In
Partnership
(a
II
non-blnding agreement
commitments)
Sustainable Development
Intergovernmental
UNESCO
2000, founded
in
2003-2004
at the
meeting
2003, extending
to
cover
(except Sudan] and
governmental
all
all
Its
fulfill
con-
World Summit on
2002.
in
to
on
It
held the
great
first
apes
at
membership during
great ape range states
key donor countries. Close
links allow
GRASP
to
operate at the
highest political levels. The partnership aims to

provide a framework into which
all
the individual
conserving the great apes and their habitats. These

Included the Jane Goodall Institutes (established

1977), the DIan
(established
In
in
Sanctuaries such as the
Fossey Gorilla Fund International
and the Orangutan Foundation
International's Care
1978],
International (established
In
1986).
Centre
Other well known International organizations
became
Fauna
including
involved,
and
Flora
International (FFI, formerly the Fauna and Flora
Preservation
the
Society),
African
in
Central
Kalimantan form part of

ttie
network for public
awareness and
Wildlife
education.
Foundation (AWF), Conservation International (CD,
and
the
International
Programme (IGCP
Captive
apes
had
In
small cages
therefore emerged.
such as the
and WWF).
transported
in
and then
conditions,
zoos or used
In
medical
International organizations
International
(IPPL), the
International
FFI,
growing animal welfare movement
research; a
League
In
AWF,
been
often
cramped and Inhumane

housed
Conservation
Gorilla
- involving
Fund
Primate
Protection
Research
staff at the
Ketambe Research
Station in Sumatra.
Ketambe
is
currently
funded by the Sumatran
Programme and
the
Born Free Foundation, and the
Leuser International
Animal Welfare (IFAW) cam-
Foundation.
for
paigned for better treatment as well as for the

conservation of great apes.
Recognizing that no one institution can solve

all
problems, partnerships have formed between
and among
NGOs and government
agencies. These
have included the International Gorilla Conservation
Programme, the Ape
Alliance,
and the Great
Apes Survival Project (GRASP) partnership. GRASP
encompasses the governments

range states, the secretariats
tional conventions,
and most
of
of the
great ape
several interna-
of the
NGOs
cerned with the study, survival, and welfare

apes.
Two United Nations agencies
Nations Environment Programme,
(the
con-
of great
United
UNEP and
the
United Nations Educational, Scientific and Cultural

Organization,
UNESCO)
lead
GRASP They
have
December
2003.
provided a joint secretariat since
Such a partnership might be expected
to last for a
243
World Atlas
and development plan-
biodiversity conservation
ning.
Such
include national biodiversity
initiatives
strategies and
action
plans,
poverty reduction
and development plans. A series
strategies,
workshops on national great ape

being held
of
each
the great ape range states, the aim of
in
each being
of
survival plans are
groundwork
to lay the
country's
participated
national
the
in
Regional Workshop
development
GRASP
also
Chimpanzee
African
\Nesl
in
for
plan.
Abidjan, Cote d'lvoire, held by
lUCN-The World Conservation Union; the national

workshop
in
Guinea; and the orangutan Population
and Habitat
Assessment Workshop
Viability
Jakarta, Indonesia;
is
it
working
in
have their recom-
to
mendations adopted by the relevant governments.

There are a number
of
other partnerships and
partnership-based projects that aim
promote
to
conservation within several great ape range states,
engage professional conserva-
including those that

Martha M. Robbins
common
tionists in
Monitoring
is
one
of the
conservation
efforts
governments,
of
essential aspects of
departments, academics, NGOs,
conservation that
others can
GRASP
is
working
to
establish through
be integrated
efficiency, effective
mobilization
and targeting
resources.
of
national great ape
recognizes the autonomy and

existing
work
in
Bwindi
are at
maximum
ensure
to
wildlife
seeks
but
initiatives,
among them.
GRASP initially appointed
GRASP
independence
of
create synergy
to
Among
the
most prominent
are:
agencies, and
communication, and successful
survival plans. Here,

field assistants
UN
action across national frontiers.
The Congo Basin Forest Partnership, which

involves
Cameroon,
Central
the
African
Republic (CAR), the Democratic Republic of

the
Congo (DRCI, Equatorial Guinea, Gabon,
and
Congo,
other nations
with
dialog
in
from outside Africa, international NGOs, and
UN
three
Special
resources and recognition
businesses.^'
aims
It
to
promote economic
Impenetrable
Envoys
National Park.
the plight of the great apes: Russell Mittermeier,
governance, and natural resource conservation
Jane Goodall, and Toshisada Nishida. They, and
by supporting a network of national parks,
to help raise
became GRASP
Special Advisor Richard Leakey,
Patrons
number
in
2003. As part of the
of
GRASP program,
missions,
seminars, and
workshops have been carried out
to help establish
of
technical
alleviation,
protected areas, and well
concessions.
related goal
improved
managed
is
to
tance to communities that depend on the conservation of the forest and wildlife resources
of 11
the current status and recent trends of each great
range states.
ape population and
The Brazzaville Process, which began
of
national policy,
their
remaining habitat;
legislation,
and conser-
vation programs; the level of law enforcement;
and
the impact on ape conservation of extractive industries
such as logging, mining, and
exploration.
oil
The plans then set out recommendations
for
forestry
channel assis-
national great ape survival plans. These identify
existing
key landscape areas
in
the participating
in
1996
with the Conference on Central African Moist

Forest Ecosystems ICEFDHAC], and involves
Burundi, Cameroon, CAR, Congo, DRC, Equatorial
Guinea, Gabon, Rwanda, and Sao
and Principe.
It
aims
Tome
to facilitate collaboration
improving existing conservation measures. Each
for the conservation
plan aims to give cohesion to the existing work of
Central African moist forest ecosystems.
many
secretariat
agencies, organizations, and individuals to
identify
is
at the
and sustainable use
lUCN Regional
of
Its
Office for
be targeted more effectively,
to
areas that are currently neglected, and
to
have been undertaken on a wide range of
envisaged
topics, including conflict resolution for forest
enable resources
to
improve opportunities for funding.
264
development, poverty
It
is
Central Africa,
in
Yaounde, Cameroon. Projects
that the plans will be integrated with other relevant
ecosystem management, sustainable use
processes and documents that relate
forest concessions, timber taxes
to
national
of
and conces-
in
play
sion fees, forest laws and policies, and critical

sites for biodiversity conservation.
A Central
Initiative
World Heritage Forest
African
ICAWHFI), wfiich
is
being developed
UNESCO and
by an alliance of
tlie
Food and
Agriculture Organization of the United Nations
regional governments,
IFAOI,
conservation NGOs, and

It
aims
respond
to
international
programs.
official aid
to tfie increasing tfireat of
hunting and the unregulated trade
illegal
in
bushmeat by promoting and supporting the
management regimes
building of
areas
in
in
transfron-
outstanding forest protected
tier clusters of
Central Africa, that will satisfy stan-
dards appropriate
status
the
to
World
of
Heritage Sites. Three such transfrontier zones
have been identified forthe
first
phase:
Gamba-
Conkouati between Gabon and Congo; Odzala-
Minkebe-Dja-Boumba-Nki between Congo,

Gabon, and Cameroon; and the Trinationale
de
Sangha between CAR, Cameroon, and
la
bushmeat
and debt
Congo.
tion
None
every
in
country that they inhabit, but enforcement has been

to
nonexistent
in
many range
states.
Even
areas designated for conservation, poaching,

logging,
and mining
ape populations.
all
in
illegal
have direct impacts on great
Typically, neither
enforcement nor
educational resources are sufficient to ensure that

conservation legislation
and abided by
in
understood, respected,
is
addition, the
In
the legislation
is
conserva-
of
of
been
these measures
easy or cheap
is
to
address the problems
and Uganda,
for instance lonce-flourishing centers
for illegal traffic in
clamped down on
chimpanzees), authorities have
this trade,
and orphaned chimp-
anzees are now rarely seen openly

countries,
for sale. In
however, agencies involved
most
wildlife
in
lack sufficient capacity, training, and resources to
undertake effective enforcement; they are some-
from committing the crime.
times also insufficiently
bushmeat have been suggested and are
great ape
being addressed
servation
NGOs
on behalf of the major con-
by the
as well as through a
initiatives.'"'
vesting
Bushmeat
number
"""'"'" '"
of
Crisis
Task Force,
intergovernmental
These ideas include
law enforcement and increased
in
increasing capacity for the
management
of
in-
fines;
pro-
result,
immune
As
to corruption.
partnerships have developed
between
government agencies and international NGOs;

these provide both financial and training resources.
International donors such as the
International
US Agency
for
Development (USAID) and the World
Bank increasingly require
rigorous,
transparent,
and independently reviewed environmental impact
assessments
(ElAsI prior to
any major development
These are seen as a mechanism
tected areas; taxing the sale of legal bushmeat;
initiatives.
promoting
courage more effective environmental protection
sources
cheap
and
of protein for
sustainable
alternative
urban consumers and rural
subsistence
hunters;
incomes
commercial traders and hunters;
for
encouraging logging and

illegal
hunting,
developing
oil
transport,
alternative
companies
to control
and consumption
of
the
Burundi
the financial benefit that the person would gain
in
Programme and
Department.
In
conservation - including the police and customs -
Various measures to control the trade
national governments have taken effective
action against poaching of great apes.
smaller than
is
confiscated
Indonesian Conservation
to
punishment specified
often a fine that
iUegally held
is
fully.
insufficient
Some
An
orangutan
by staff of the Sumatran
implement, and action on the ground has so far
Great apes are protected by national law
poor
measures
performance.
NATIONAL AND INTERNATIONAL LAW

National legislation and enforcement
concessions; and linking of aid
at their
relief to verifiable
developing countries. EIA legislation

poorly developed and/or policed
in
is
many
to enin
however
great ape
range states. Where such assessments are conducted, they tend to focus on site-specific impacts
rather than the wider socioeconomic effects that
265
World Atlas
Convention on Biological Diversity
The
CBD came
on December 29 1993.
into force
It
establishes three main goals: the conservation of

biological
diversity,
the
sustainable use of
its
fair
and equitable sharing
of
components, and the
the benefits arising from their use. Although

not
list
does
it
species or places of particular conservation
concern, the themes, principles, and activities of

the
It
CBD
ways
border.
orphans destined
the conventions to find
to
promote transfrontier co-
to
communities
local
ecosystem management.
in
Thematic programs covering the biodiversity
Many
Inland waters,
for
conservation of the habitat of
are confiscated and
example
placed
CBD on
in
sanctuaries;
logging
eventually return
Volker
might lead, for example,
an increase
to
of
ElAs
or around ape habitats,
in
but proper enforcement
is
to
have any value. For an EIA
needs
to
be commissioned with
measures
must then be implemented. Plans
up industrial logging
of the civil
to
much
in
of
DRC
that
to
it
and
it
open
following the
war, and backed by the World Bank,
suggest that there
ceases
be useful,
to
a serious intent
the mitigation and harm-avoidance

specifies
mea-
required for such
sures
end
need
is
improve the regulations on the content and
implementation
is still
be acceptable
a long
way
to
go before
it
to liquidate a nation's forest
estate and biodiversity resources
in
An
the Dja Biosphere Reserve
home
is
western lowland
In
forest
of
Cameroon,
chimpanzees and
to central
gorillas.'''"
bushmeat
in
hunting, trade, and consumption. There

to
Sommer
the great apes.
impact and management
the
an area that
to the wild.
all
the case study report produced for the
Is
In
of
and mountains address
forests,
export or internal trade
some
concerns during
biodiversity
operation, and to involve indigenous peoples and
Ibadan Zoo, Nigeria,
was captured near the

Cameroon
deal with
to
development planning,
This young chimpanzee

in
are very relevant to great ape conservation.
encourages parties
the
name
of
in
Endangered
Species of Wild Fauna and Flora
Awareness
of the threat that
unsustainable Inter-
many animal and plant

which was signed in 1973 and
national trade posed to
species led to CITES,
came
Into force in
national trade
1975. By regulating the inter-
endangered species
In
las listed
in its
appendices], CITES aims to ensure that Interna-
specimens ofwild animals and plants
tional trade In
does not threaten their
CITES
survival. Parties to
are expected to implement the convention through
CITES accords varying
their national legislation.
degrees
protection
of
to
approximately 33 000
species of animals and plants, whether they are
national development.'"
traded alive or as "readily recognizable parts or

derivatives' such as dried or
Membership
souvenirs, trophies, tusks, or timber All the great
A number
of international conventions
of international
conventions that address
different aspects of biodiversity conservation
been agreed and are

vance
to great
Trade
In
in
have
force. Of particular rele-
apes are the Convention on Biological
Diversity (CBDl,
the Convention on
Endangered Species
of
International
Wild Fauna and
apes are threatened by hunting for food,

curlos; although
much
domestic,
High levels
of
much
of
or as
pets,
and
the resulting trade
chimpanzee trade occurred during
the 1950s and 1960s; at least 300 infant chimpan-
zees were reported
to
have been exported
to
Europe
during 1950-1956 alone, with about 100 deaths
the Convention on the Conservation of Migratory
transit.^
of
Wild Animals (CMSl, and a
number
of
regional conventions. The extent to which the range

states are parties to these conventions
marized
in
Table 14.1.
is
sum-
Is
also crosses international borders.
Flora (CITES], the World Heritage Convention IWHCI,
Species
U6
smoked meat,
International conventions and compliance
Much
of this trade
was
driven by
chimpanzees
In
demand
use
In
species of great apes are listed
in
from the market
for live
for
biomedical research."
All
Appendix
six
I
of
CITES, which
means
that international
trade
them
in
permitted only
is
exceptional
in
circumstances, and never primarily for commercial
A permit
purposes.
may be
any
to trade
apes
of the great
movement
Europe, North America, and Asia.
A systematic
uation of the accuracy of CITES reporting
in
eval-
relation to
specimen
the overall great ape trade has not been undertaken,
not detrimental to the survival of the species, as
although widespread lack of CITES compliance has
in
the exchange of breeding individuals between
the
if
of the
reputable zoos. The relevant country's designated
proposed recipient
and care
for
that has
been
All
must be
authority
scientific
any traded
listed in
live
Appendix
international trade
great ape
U6
involved
specimens
(831 of
of all
Much
reintroduction
P.
apes
is
in
en-
dangered species, CITES also addresses a number

conservation of
species affected by trade. One such issue
bushmeat
trade; this
CITES
be an issue
to
is
is
the
considered by the parties to

of
both trade and wildlife
of
ACC
Range
CBD''
CITES'
CMS'
WHC"
members
Angola
yes'
no
no
yes
no
Burundi
yes
yes
no
yes
signed
Cameroon
yes
yes
yes
yes
yes
yes
yes
no
yes
yes
Congo
yes
yes
yes
yes
yes
Cole d'lvoire
yes
yes
yes
yes
yes
Democratic Republic
yes
yes
yes
yes
yes
no
of
programs, for
and
the wild,
for
scientific
of this trade 119-23 percent]
move across
Table H.I Great ape range states: parties to international conventions
paniscus 112
accounted for by great apes that are part

that regularly
171
between zoos and
captive-breeding
into
Some
purposes.
their
which were captive bred].
of
of the trade in live
for
the recent past."
in
addition to the regulation of trade
In
species
subspecies
which were captive bred), 30
sanctuaries,
from Indonesia
28^ Pan troglodytes
which were captive bred), and 324
Pongo species 1249
the African range states,^' and
between 1975 and 2003
reported
Gorilla of several
which were captive bred),
of
must report
of CITES-listed species. Total
in live
in
of issues that are pertinent to the
I.
great ape range states apart from Angola
imports and exports
been documented
orangutans have been traded under false papers
house
animal or plant species
are parties to CITES, and so
of
the
that
satisfied
suitably equipped to
is
puy
African nations or from great ape range states to
is
issued only
in
was
circuses
of
international borders,
of the
state
Congo
Equatorial Guinea
yes
yes
no
no
sometimes over-
Gabon
yes
yes
no
yes
no
some
Ghana
yes
yes
yes
yes
yes
moved between
Guinea
yes
yes
yes
yes
signed
Guinea-Bissau
yes
yes
yes
no
no
Indonesia
yes
yes
no
yes
Liberia
yes
yes
yes
yes
yes
Malaysia
yes
yes
no
yes
Mall
yes
yes
yes
yes
yes
Nigeria
yes
yes
yes
yes
yes
Rwanda
yes
yes
yes
yes
yes
Senegal
yes
yes
yes
yes
yes
have been traded between 1975 and 2003 for this
Sierra Leone
yes
yes
no
yes
signed
purpose. The source Iwild or captive bred) of 37
Sudan
yes
yes
no
yes
yes
United Republic
yes
yes
yes
yes
yes
yes
yes
yes
yes
yes
most
of
whom were
bred
noted that these trade
in
captivity.
data
estimate the volume of the trade as,

instances, an animal that has been
countries on
more than one occasion
must be
It
will
in
generate
multiple records over the course of a year
As well as
live
body parts, blood,

entific
individuals, trade
hair,
use. Of the
can include
or other specimens for sci-
specimens reported
in
inter-
national trade for biomedical purposes, the majority

involved
57
live
samples
of blood, hair, skin etc.; a total of
chimpanzees were, however, reported
these animals
was
not reported; the others
to
of
were
from captive sources, or animals obtained prior to

the listing of the species on CITES, or from a range
state that
was
not yet party to CITES.
Some
trade
of
Uganda
in
a
the bodies of great apes has also been reported.
The bodies
Tanzania
CBD, Convention on
Biological Diversity.
b CITES, Convention on International Trade
were exported as hunting
CMS, Convention on
the United States of
WHC, World
America and France during 1995. These may have
ACC, African Convention on
yes: the country
of five gorillas
trophies from
the
CAR
to
been trophy animals obtained prior
to the listing of
the species on CITES.
Most trade
in
wild
is
either
Conservation
of
in
Endangered Species
of
Wild Fauna and Flora,
Migratory Species of Wild Animals.
Heritage Convention.
is
tfie
Conservation of Nature and Natural Resources.
a party to the convention;
signed: the country has signed but

no: the country
apes
ttie
is
not yet a
full party;
has not signed.
between
247
"
World Atlas
management. Where cross-border trade

meat occurs,
Consequently,
range and donor states.

raised by the trade
It
aims
to
bushmeat,
in
examine issues
to identify solu-
promote awareness and action
to
of the
bushmeat
The group's
trade.
to
management
work was
achieve better and more sustainable
it
may be
initial
on a case-study area comprising Cameroon, CAR,
in
200i
conservation and trade

with
GRASP and
the
in
work
great apes, and to
in
CBD
on progress
to report
The
of great
WHC
areas that are
live in
rich,
or particularly
noteworthy for other reasons. The importance
some
of
these has been recognized by the
which aims
cultural
to
of
WHC,
define and conserve the worlds
and natural heritage. The
adopted by the General Conference
of
WHC
was
UNESCO
1972 and has compiled the World Heritage
in
List,
comprising sites or areas nominated according
to
Criterion
The forest edge
at
and
livl
servation
National Park, one of the
containing
under the convention's
should contain "the most important
significant
Bwindi Impenetrable
natural habitats for in-situ con-
biological
diversity,
threatened
species
of
including
of
those
outstanding
national parl<s that has
universal value from the point of view of science or
World Heritage
conservation."
status.
If
World Heritage
must report on
the condition of the sites within their borders, on
measures taken
conserve them, and on efforts
to
to
awareness about them. Of the 23 great
raise public
ape range states, eight have World Heritage Sites
harbor
that
populations
of
apes
great
Isee
Table U.2I.
the
July 200A.
In
Park Isee Box
Sumatra was designated
.2) in
part
Cluster Mountain' World Heritage Site; this
of a
Kerinci
Site is threatened
neither containing
Seblat,
This initiative derived from an
government
earlier proposal from the Indonesian
UNESCO award World
request that
to
Heritage Site
status to the whole Leuser Ecosystem of about
26 000 km', rather than to only the 8 900 km^ of

the
for this
was
Ecosystem
in
3 573
estimated
(an
The
Park.'"'^
most orangutans
that
rationale
the Leuser
out
of
598
individuals! live outside the national park, so
an
excessive focus on the park alone would risk major
both
losses of
specified criteria.
Areas nominated
has been signed by more than 175
orangutans!.
apes
species
in
includes two other parks to the south [Bukit Barisan
Secretariat.
World Heritage Convention
Many populations
World Heritage
Parties to the convention
states.
Selatan and
highly distinctive,
of
intended to draw the matter to the
is
world's attention.
Congo, DRC, Equatorial Guinea, and Gabon. CITES
adopted a resolution
placed on the List
Danger, which
of interested
implemented by range
tions that can willingly be
and
bush-
2000, the parties to CITES set up a
in
Bushmeat Working Group, composed
states,
in
often unsustainable and illegal.
IS
it
and orangutans. Cluster
habitat
Mountain World Heritage Sites are

in
typically located
high mountain ranges Isuch as the Himalayas,
Andes, or Alps!, where the focus

is
all
conserva-
considered this designation to be appro-
tionists
priate
conservation
of
on the high-altitude habitats. Not
for
use
in
biodiversity occurs
Sumatra, where the greatest

in,
and
is totally
dependent
on,
lowland forests.
Gordon
IvIiHer/IRF
World Heritage Sites are also protected by the

highest category of national designation, normally
as a
national park';
apes
live
World Heritage
that are
many
of
those
in
which great
have, however, been placed on the List of
home
in
to
Danger.
In
particular, sites
in
DRC
eastern gorillas, chimpanzees, and
bonobos have been seriously affected by armed

conflict
and
its
consequences. The World Heritage
Committee has therefore undertaken

support and assistance to
lUCN and other
DRC
institutions,
in
to
provide
cooperation with
such as the World Bank
and the United Nations Foundation.

It
of
has been proposed that a new designation
World Heritage Species be created, and that
all
the great apes be given this status." This idea enjoys
strong,
although not universal, support
conservationists.'^^
2/i8
If
this
among
proposal were accepted,
Table
U.2 World Heritage
Range
state
play
Sites containing great ape populations
World Heritage
Year
Species or
inscribed
subspecies
Site
World Heritage
in Danger
Cameroon
Dja Faunal Reserve
Cote d'lvoire
Tai
Democratic Republic
Virunga National Park
1979
mountain gorilla.
chimpanzee
Kahuzi-Biega National Park
1980
eastern lowland
(/
/
/
^1
1987
western lowland
qonlla,
National Park
Comoe
National Park
chimpanzee
chimpanzee
chimpanzee
1982
1983
Congo
of the
in
Salonga National Park
1984
Okapi Faunal Reserve
1996
chimpanzee
chimpanzee
bonobo
chimpanzee
Mount Nimba
1981
chimpanzee
2004
Sumatran orangutan
gorilla,
Garamba
Guinea and Cote
d'lvoire
1980
Nature Reserve
Strict
Indonesia
National Park
Tropical Rainforest
Heritage of Sumatra
Malaysia
Kinabalu Park
2000
Bornean orangutan
Senegal
Niokolo-Koba National Park
1981
chimpanzee
Uganda
Bw/indi
Impenetrable
1994
mountain gorilla,
chimpanzee
chimpanzee
National Park
Rwenzon Mountains
1994
National Park
new World Heritage
Sites
would then be designated
specifically for the protection of viable populations

of
A new
these species.
Protocol to the World
Heritage Convention would be needed to establish

this
new
category.
three of the mountain gorilla range states,
All
IDRC, Rwanda, and Uganda! are parties
Rwanda was
the
most recent
The Convention Secretariat has agreed

a possible
to
agreement, under Article
CMS.
to
June 2005.
to join, in
promote
to
iV of
CMS,
the
cover the conservation of the shared mountain
Convention on the Conservation of Migratory
gorilla
Species
agreement could also cover key World Heritage and
The
CMS aims
to
conserve
and avian migratory species
all
dictions that they visit or through
Appendix
of the
Ct^S
lists
marine,
terrestrial,
in
migratory species that,
according to the best scientific evidence available,

are endangered. Although not migratory
sense,
mountain
gorillas
other protected sites."
national juris-
which they pass.
in
population of the three countries. This
nation
the Terrestrial
2004,
In
Group
of
CMS
the
mountain
of
Mammals Working
proposed that the special desiggorillas
be extended
inter-
Compliance with international agreements
national borders and, because they are of particular
The success
conservation concern, the subspecies
the ability of the parties to implement
Appendix
gorilla
of the
CMS. Since
has also been designated for
problems facing
its
habitat.
listed
in
1997, the mountain
Action' on the basis of the conflicts
in
This
Concerted
DRC and
the
designation
requires parties to undertake particular actions to
help conserve the subspecies and

Scientific Council of the
CMS
its
habitat.
The
has also noted the
opportunities for cooperation with other conventions

in
all
the usual
regularly cross
is
to
gorillas.
protecting the habitat of mountain gorillas
in
DRC.
of all
fectively, including
reporting
mand
to
these conventions depends on

ef-
requirements. These commitments de-
the allocation of trained staff and resources
cover operating costs;
priorities are
the
them
compliance with the conventions'
means
when other government
taken into account this
of
is
often beyond
great ape range states.
prehensive assessment
No com-
of the effectiveness of
conventions has yet been carried out, but the
has adopted a measurable
target.''^
This
is
these
CBD
the '2010
2M
World Atlas
on Africa Forest Law Enforcement and Governance

lAFLEGl.'"
The Yaounde Forest Summit
1999 brought
of
together leaders from the Congo Basin countries

to
address the growing threats
region and to look at
and
integrity
ways
help ensure their
The principal outcome was
survival.
Yaounde Declaration'
the
to the forests of the
to
intent
of
to
promote
transfrontier cooperation for biodiversity conservation
and around the Congo Basin,
in
involving
Cameroon, CAR, Chad, Congo, DRC, Equatorial

Guinea, and Gabon.
It
established firm political
commitments, such as the creation
of
new
forest
protected areas, developed plans to combat illegal

logging and poaching of wildlife, and broadened
the application of sustainable forest-management
Gordon Miller/IRF
billboard at the
target', by
which the parties committed themselves

2010 a significant reduction
strategies. This paved the
new
Cameroon, Congo, and CAR
Equator crossing near
to achieve by
Lake Mburo National
rent rate of biodiversity loss at the global, regional,
ed collaborative
and national
of forest with
Park, Uganda,
warns
of
of the cur-
level.
the dangers of
corruption.
way
and
for both action
partnerships, such as the accord
between
2000, that establish-
in
management
over 28 000 km'
of
harmonized forestry
newly created Trinationale de
policies, in the
Sangha conser-
la
Regional agreements and activities
vation area. To date, a total of 34 000 km"" of
Regional agreements
protected areas have either already been gazetted
apes are
All African
Class
under
listed
of the African
'full
protection'
Convention on the Conser-
vation of Nature and Natural Resources (ACCl. This
entered into force
in
1969, and
is
binding on the vast
or are
able
It
IS
designed
to validate
and encourage
been made
management
amount
to a total
to
in
Congo Basin;
the
strengthen the sustain-
of existing protected areas,
Conference on the
Ministerial
Environment, held
in
Maputo
which
kml'"
area of 135 000
The African
majority of Africa's 21 great ape range states liable

14.11."''
the process of being so
in
efforts have
new
in
2003, adopted the
conservation and the wise use of natural resources
Action Plan of the Environmental Initiative lAPEI]
by individual countries and by countries working
of the
together
significance
Its
and a basis
for
is
mainly as a policy guide
developing specific measures to
conserve resources such as great apes that are

important "from an economic, nutritional,
scientific,
educational, cultural and aesthetic point of view,"
in
the words of the convention.
is
an
initiative
organized by donors
such as the World Bank, and involves the

pation of governments,
NGOs, and
civil
focuses on combating the threat posed

by
logging
illegal
Partnership for African Development
Initiative
to
[APAII,
which
is
an Africa-led
partici-
society
It
to forests
and trade, corruption, and
initiative
mobilize African institutions and expertise to
enhance the
role of protected
areas as tools for
safeguarding biodiversity, sustaining ecosystem

processes, and contributing to
The Forest Law Enforcement and Governance

IFLEGI process
New
[NEPADI. This includes the African Protected Areas
livelihoods
and
sustainable development across the continent.
aims
to
encourage and enable the development
It
of
conservation areas, the building of the capacity to

both implement the
CBD and to manage
and knowledge about
it,
biodiversity
and the networking
of
African experts as well as institutions.
poaching. Ministerial and other meetings have been
organized
to
in
(FLEG
Asia),
Regional and species action plans
A number
of regional action plans,
which involve
cers and consumers, the private and public sector,
several range states, and species action plans have
and donors, that can help address
been developed
exploitation.
250
Africa (AFLEGl and Asia
examine potential partnerships between produ-
These meetings have
illegal forest
led to ministerial
of great
to
guide and organize conservation
apes. Notable recent examples from the
declarations and action plans, an example being
Primate Specialist Group
the October 2003 Yaounde Ministerial Declaration
Survival
Commission ISSCl
of
the
include:
lUCN Species
Many
now
partners
Status survey and conservation action plan for
apes.
African primates;'"'
(see Annexl, and their activities are
Status survey and conservation action plan for
West African cfiimpanzees;
"
"' "
IPHVAI for mountain gorillas:
chimpanzees
for eastern
volume, especially
Chapters 16 and
Population and habitat viability assessment
PHVA
in this
these are
of
in
touched on here
17.
in
GRASP
in
summarized
in
Their activities can only be
to give
an idea
of the
The international NGOs
Uganda;"
breadth
of
active in Africa include
the International Gorilla Conservation

for
play
their involvement.
and
PHVA
in
orangutans."'""
(IGCPI
partnership
(as
Programme
discussed above
in
the
section entitled 'Partnerships'), Conservation Inter-
A number
of
other species-specific plans have also
been developed, including two on bonobos,"'

one on chimpanzees
ments are sets
of
into
and
Uganda.'" These docu-
in
recommendations; prior
involvement of GRASP, few appear

incorporated
'"
documents
national planning
WCS
jects
it
The lUCNs West African Chimpanzee Action

Plan reviews existing information on the status of
the two subspecies that occur
West
in
Africa (up
border between Nigeria and Cameroon), Pan
to the
comprises
troglodytes verusand P.
t.
a set of national profiles
and a regional assessment
of
veiterosus.'^
recommendations
threats and
It
for
action.
It
Fauna and Flora International

Conservation
(FFI),
(WCS),
Society
and
Global Conservation Organization.
has a very active program
in
Central and
East Africa, including great ape conservation pro-
such as national biodiversity strategies and action

plans.
(CI).
Wildlife
WWF-The
to the
have been
to
national
the
in
Congo, Gabon, DRC, Uganda, and Rwanda.
CI
has a major program
has identified as
in
West Africa,
a region
a 'hotspof for biodiversity."
It
also secured core funding and provided technical
and
logistical
support for the development
of the
lUCN's West African Chimpanzee Action Plan.

IGCP's goal
is
the conservation of mountain gorillas
and their habitats
and
to
in
DRC, Uganda, and Rwanda,
increase cooperation between their pro-
tected area authorities.
Despite
civil
unrest and
assesses logging, agriculture, crop raiding, bush-
other setbacks, the project has achieved consid-
meat hunting, primate sanctuaries,
erable success
rehabilitation
experiences, and the threat from infectious diseases.
The plan was based on

in
workshop held
in
2002
Abidjan, Cote d'lvoire, attended by 72 scientists,
conservationists,
makers, protected area
policy
As well as
are facilitated by the lUCN's Conser-
helping the authorities protect
in
gorillas
in
its
the Virungas since the 1980s.
role in IGCP,
WWF
has projects
across Africa, including support for forest conservation projects
and CAR.
managers, and donors from 15 countries."
PHVAs
mountain
in
Cameroon, Cote
d'lvoire,
Gabon,
rangers' truck sports
the
WWF logo. WWF was
one
of the first
become
NGOs to
involved in great
ape conservation issues.
Gordon Miller/IRF
vation Breeding Specialist Group, but involve local
stakeholders and various organizations with an

interest in the conservation of target species.
The
process uses mathematical models within a participatory
workshop framework
to
produce a strategic
recovery plan for a threatened
species and
its
The process uses data on demography,
habitat.
genetics, and ecology, as well as estimates of threats
such as current and predicted land-use patterns;

it
is
explicitly
designed
to
broaden stakeholder
involvement and enhance knowledge sharing."'"
PHVA workshop
held
in
that focused on orangutans
Jakarta, Indonesia,
in
was
January 200^.
NONGOVERNMENTAL ORGANIZATIONS
A
large
number
working
local
in
of international
and national NGOs,
partnership with national authorities and
NGOs, have been
involved
in
developing and
implementing conservation programs
for the great
251
World Atlas
other groups active
the field include Les
in
Amis des Bonobos du Congo, BerggoriUa und

Regenwald
the
Direkthilfe,
Bonobo Conservation
Born Free Foundation, Cameroon Wildlife
Initiative.
Aid Fund, Dian Fossey Gorilla Fund Europe, Dian
Fossey
Fund
Gorilla
Frankfurt
International,
Zoological Society, the Jane Goodall Institutes, Pan
Chimpanzee
African Sanctuary Alliance, the Wild
Foundation, and the Zoological Society of London.

In
in field
Southeast Asia,
much NGO
conservation and orangutan rehabilitation.
Malaysian Borneo,
In
the
Kinabatangan River, Sabah.
Borneo, the Orangutan
funds patrols
National
Park,
and releases orphan orangutans
rehabilitates
Lamandau
Indonesian
Foundation International
Tanjung Puting
in
project along
In
Strict
in
Nature Reserve, and supports
research into conservation and forest restoration.
The
Borneo
rehabilitates
Orangutan
and releases orphans
and other parts

proposals
of
in
Kalimantan, and
to protect the
peat-swamp
Survival
in
Mawas
to 30.
The LDP
year phase.'^ On the other hand,
in its first
early 200^ the
in
which was planned
Sabah,
In
pass through the
to
Gunung Leuser National Park
Box
(but see
Kinabatangan
the
11.21.
Orangutan
Conservation Project IKOCPI was established
1998
secure one
to
populations,
in
the
of
largest
Kinabatangan floodplain
the
KOCP
eastern Sabah. The
Malaysia and works closely with the Sabah Wildlife
Department,
holders
communities, and other stake-
local
to find
ways
conserve wild orangutans
to
orangutan population
in
uals
km'
of
in
of
KOCP
several thousand individ-
the Kalabakan area, between the Maliau
Danum
Basin and
Valley Conservation Areas." This
was assessed as being perhaps
the population of
bility,
is
in
Jambi province.
also the only organization that
carrying out field research
research
site
open;
it
in
is
currently
Ketambe, keeping the
conducts most
of the
sun/ey
provided that logging Iboth legal and
be brought under control and associated
forest
fires
escape
agreement
terminated
workers on
when
to convert 2
distribution of the wild orangutan population in the
related networks exist, linking
prevented. The area
2002,
in
into
Two
been used as an excuse

of all the trees, with a
the Orangutan Conservation Forum, proposed by
PHVA workshop, which aims
the 200^
including
advising on the preparation of the national great ape

survival plans for Indonesia and Malaysia.
habitat for
Sumatran orangutans.
Gunung Leuser National Park
It
18
the provinces of North
in total.
The whole area
is
to
a key
includes the
900 km'l and
about 26 000 km'
the subject of the long-
term Leuser Development Programme (LDPl, a

partnership
project
was
for clearfelling (the
view
removal
subsequent planting]
to
of forest.
PROTECTED AREAS
Overview
In
is
Sumatra and Nanggroe
Aceh Darussalam, amounting
from extensive areas
'^^
11.21
extensive areas of protection and production forests

in
natural forest
plantation
to coordinate
education and communication efforts,
The Leuser Ecosystem (see Box
of
existence of this agreement, however, had already
research scientists involved
conservation;"' and
4U km'
Sabah government.'"' The
the
by
had a narrow
Malaysian-Chinese
mangium pulpwood
Acacia
both islands; the Orangutan Network, which links

in
via-
illegal)
could
and monitoring work concerning the status and
area.
in
has identified an
ISOCPI has established a similar release program

Bukit Tiga Puluh National Park
of
WWF-
supported by
is
in
Malaysian
Bornean orangutans with the best long-term
It
LIF
nine
go ahead for the Ladia Galaska road
initial
project,
involved
The
created
Indonesian president Megawati Sukarnoputri gave
Balikpapan
area, 5 000
NGO
now supporting
is
managing the Leuser Ecosystem
multiple-use forests. The
forest inhabited by orangutans.
an
Sumatran Orangutan Conservation Programme
in
between the government
of
terms
of their habitat
great apes and
humans
needs, the interests of
often
come
into conflict;
on
the whole, apes need natural forests free of distur-
bance while humans
rely
on farmland, plantations,
or forests that are intensively
managed
for pro-
duction purposes. Areas of great ape habitat must

therefore be set aside with permanent legal and
actual protection
the wild. The
if
ape populations are
same
analysis applies to
to survive in
many
other
tropical moist forest organisms, so protected areas
means
Indonesia and the European Union, on which the
are the chief
EU has spent around US$39
type of ecosystem.'" National parks and other areas
million
and the
Indonesian government about US$7.5 million since
252
extendable
Foundation
is
to
Leuser International Found-
ation (LIFl, which has these rights for nine years,
Hutan runs an innovative
community conservation and research
whole area
of the
for this purpose, the
the
effort is invested
The Indonesian government has assigned
1996.
management
where the
forest
of
may
conserving biodiversity
not be
damaged and
in this
wildlife
may
in
play
not be hunted are fundamental to the con-
servation of great apes. All great ape range states
have
systems, typically covering
protected area
5-15 percent
relatively
few
of
these areas are important for great
apes and some,
in reality,
exist only
on paper.
particularly challenging for the smaller
is
It
although
of their national territory,
areas
protected
maintain
to
ape
healthy
population. Small populations of great apes are

highly susceptible
due
extinction
to
random
to
catastrophes (such as infection by Ebola virus, or
exposure
reduced genetic
to forest fires],
human
poaching, and other forms of
Moreover, the
nities
variability,
disturbance.
commu-
transfer between
ability to
essential for the viability of great ape
is
populations; one that
composed
is
of
small isolated
groups has only a precarious future. Population

densities of great apes are typically only 0.3-1.Q
per square
individual
kilometer;
areas
large
of
protected habitat are therefore needed to support
populations that are
be viable
likely to
in
term. Such populations should generally be

least
000 individuals
hundred
to
Great apes
in existing
protected areas
species, includes within
and unprotected
Tanjung
of at
of several
several thousand square kilometers.'"
The Bornean orangutan,
containing
an area
living in
any widespread
like
its
range both protected
forests. Indonesian national
parks
orangutan populations include Kutai,
Gunung Palung, and
Puting,
Betung
Kerihun. Most Sumatran orangutans are found
Leuser Ecosystem, which
the
Michael Huffman
the long
management
but
still
is
in
under active
subject to illegal logging.
In
Malaysian Borneo, the most significant population

in
Sarawak
is
found
in
the Lanjak-Entimau Wildlife
Sanctuary [which now incorporates Batang

National Park and
In
is
connected
Ai
Betung Kerihun).
to
Malaysia, a wildlife sanctuary
is
more
strictly
protected under state legislation than a national

park. Likewise, orangutans occur
in
several areas
Sabah that are strongly protected under

legislation, including the
Danum
in
state
Valley Conservation
Area.
have yet
be surveyed. Similarly, no precise
to
Top:
estimate exists for protected area coverage for the

central
chimpanzee or western lowland
of the existing population of the
lowland gorillas
is
within formally protected areas
(although the status of this population

to
reported
eastern
is
uncertain
heavy poaching, and insecurity
preventing surveys), and
all
mountain
gorillas in the
Virungas and Bwindi are within national parks. The

position for the eastern
because extensive areas
chimpanzee
of the
is
less clear
eastern Congo Basin
percent of
the
its
distribution),
and three
in
6.6
the range of
Nigeria-Cameroon chimpanzee with similar
coverage."
cially
Finally,
bonobos occur
protected areas
in
358 km'].
chimpanzees were
released into the wild
in
the 1960s, growing by
2002
to a self-sustaining
only two offi-
population of 60.
DRC. one very large
Above: Deforestation up
in
(Salonga National Park, 36 560 km^) and the other
much smaller
Island,
Tanzania, v\(here 17
gorilla.
chimpanzee's geographical range (covering
Rubondo
United Republic of
There are 26 protected areas within the western
Much
due
Martfia M. Robbins
(Luo Reserve for Scientific Research,
to the
boundary
of
Bwindi Impenetrable
253
World Atlas
The effectiveness
of protected areas
As with law enforcement, nnany
ment programs
the benefits of conservation, and there
existing nnanage-
for protected areas
in
Southeast Asia are heavily funded by
Africa
and
NGOs and
some
In
local
cases.
communities and
It
may be
possible
without this support. Iâny of the national parks
tourism,
and reserves
(from documentary makers), and local community
great
in
ape range states are
understaffed and underfunded, and are
more than 'paper
parks'.'" Indeed, by 2000,
declining great ape populations had been reported

in
almost
all of
2U protected areas where great ape
meet both requirements through Income from

research
areas
in
the
in
collapse of eastern
functional
990s has occurred despite the large
not realistic to expect national
is
It
bear
to
all
protected
the costs of maintaining
areas.
example, the opportunity cost
obtained from using a resource

talnably or otherwise, because
better
in
the long term than areas that have no
protected status at
all,
as people often respect
their boundaries. ''"They also deter certain kinds

of
Investments,
such as major infrastructure
to the
government
of not
has been estimated

annually;"'
a 2 700
In this
that the International
mental impact assessments, and by public opinion
It
some
notice of them.
at
beneficial by local people as
National Park, Rwanda,
well as by governments
destroying snares set
maintained. Park administrations therefore usually
by poachers.
try to
If
Governments
the value
least
will naturally
argue
community should contribute

the global environmental
of
benefits generated by such an Investment.
has long been accepted that protected areas
must be perceived as
being protected)
logging a tropical forest
km* protected area would exceed US$40
which are bound by law
to take
other ways, sus-
Is
about US$15 000 per km^
at
million per year
commission environ-
In
It
for
the revenues not
sense, the cost of setting aside
projects by publicly accountable donor agencies,

to
Cameroon,
In
(I.e.
protected areas overlapping their range. On the
generally fare
all
'" For
Central Africa
other hand, even
'paper parks'
fees
filming
however, feasible for
this Is not,
Congo Basin."
particular.
governments
lowland gorilla populations
communities,
development projects, supplemented by external

donor support;
research programs had been conducted.' The
since the late
Volcanoes
governments.
Increasing
to
little
in
between
that can be shared
is
generate revenues
to
external donors and would not be able to function
critically
Rangers
pressure for protected areas
they are to survive and be
reach out to nearby communities
to explain
This
policy
Facility
'incremental' cost of
basis
(GEFj,
in
price
Global
the
for
which finances the
measures aimed
conservation. This Incremental cost
Is
at biodiversity
the difference
between what a responsible government
should Invest
Ian
the
Is
Environment
the Interests of
In
Its
own people
le.g.
Redmond/UNESCO
to
secure water catchment services and tourism
revenues], and what the world
have done
like to
community would
secure a globally Important
le.g. to
ecosystem, or the carbon-storage function

forests). Part of the
slow,
is
that conservation
of
is
long-term business, requiring recurrent
Investment
donor
challenge
in
interest
perpetuity, while
in
governmental and
any given conservation
initiative
operates within shorter timeframes. To help solve

this,
the
GEF has
often
endowed
trust funds
mechanisms, equipping them
trust-like
to
and
invest
a one-time-only grant with the intention that the
return on capital will underwrite a share of conservation
expenditure
was used
In
indefinitely."
the Mgahinga-Bwindi
method
This
Impenetrable
Forest Conservation Trust, centered on two national
parks
illas
In
southwest Uganda where mountain gor-
occur'^' This trust
Initial
was
funds from the GEF;
source
of
established
it
in
995 with
funding for park management, research,
and community conservation projects;'"

such examples are
254
provides a sustainable
rare.
globally,
in
play
TRANSFRONTIER PROTECTED AREAS

Many
great ape populations extend across national
these coincide with protected
frontiers; in places,
lor
otherwise managed) habitat areas that have
been established on either side

principle,
managers
in
of the best
mountain
border.
In
neighboring countries to cooperate to
improve great ape protection
One
of
these provide opportunities for reserve
examples
habitat
gorilla
management
is
in
combined
the Virungas,
in
which the authorities
of
area.
the cluster of protected
Uganda, and the DRC cooperate
Yaounde Ministerial Declaration
of
the
in
Rwanda,
closely.
The
1999
was
of
intended to promote transfrontier cooperation for
and around the Congo
biodiversity conservation in
Basin.
Among
other things, so far
recognition and
endorsement
Trinationale de
of
it
has
led to:
of the creation
Sangha, which has a
la
core area of around 7 300 km',
(Lobeke National Park),
in
Cameroon
CAR (Dzanga-Ndoki
National Park), and Congo INouabale-Ndoki

National Park);
new
the creation of two
forest national parks
Cameroon ICampo Maan and Mbam
in
et
Gordon Milter/IRF
incorporates the Lanjak-Entimau Wildlife Sanctuary

in
Sarawak and the Betung Kerihun National Park
in
West Kalimantan, Indonesia, a complex with a
combined
total area of 11
being coordinated
in
adjacent to Virunga
National Park
In
the
and the Forest Depart-
the Congo and to the
Park Management Unit

Park
Is
(ITTO)
ment
between Congo IBambama-
initiative is
Sarawak, with WWF-lndonesia and the
Timber Organization
Lekana National Park! and Gabon IBateke
This
Gorilla
Uganda
International Tropical
DjeremI;
vation initiative
kml
In
by the
the establishment of a transborder conser-
in
000
Mgahlnga
National Park
of
Betung Kerihun National
In
Rwanda.
Indonesia.
Plateau National Park);

the founding
in
2002
of the
Congo Basin Forest
Partnership, which seeks to build upon

existing unilateral, bilateral,
programs
tion
many
and multilateral
the region, including collabora-
in
between governmental and
NGO
stake-
MANAGEMENT OUTSIDE PROTECTED AREAS

Logging
Most great apes
of great
ape habitat
consider
A program funded
by the European Union (Appui
Gestion Integree des Ressources Naturelles
des bassins du Haut Niger et de
AGIRl
is
working
in
West
la
Haute Gambie,
Africa
to
establish
transboundary conservation agreements between

Guinea and neighboring countries with contiguous
forest,
much
includes
in
of
which
is
chimpanzee
habitat.
It
Niokola-Koba National Park ISenegall;
Badiar National Park (Guinea); Mount
Nimba
Strict
how
management
the wider environment
crucial to their survival.
holders.
a la
outside existing or planned
live
protected areas,'" so protection and
We
is
should therefore
the survival of great apes can be
reconciled with logging, which
is
one
of the
major
influences on their habitats outside protected areas.
To do
this,
it
is
necessary to understand more
fully
the impact of timber harvesting on great ape behavior and ecology.
For gorillas, selective logging

availability
of
digestible
easily
similar plant foods
the
their environment. For these
in
animals, the threat
may improve
herbaceous and
is
less the alteration to the
ecosystem caused by logging, but rather the
Nature Reserve IGuinea-Cote dTvoire); the pro-
forest
posed Bafing-Faleme Protected Area (Guinea-
greatly increased hunting pressure brought about
and the proposed Guinea-Bissau-Guinea
by forestry workers living off the land, and the
Mali);
Protected Area.
Since 1995, there have been efforts to establish

a transfrontier biodiversity conservation area that
improved
access
to
commercial hunters and
transport opportunities provided by logging roads
and vehicles. The situation for orangutans
is
less
255
World Atlas
much
taking
greater care
process to preserve
The
latter
in
logging
the
fruit trees.
would include:
taking a detailed inventory prior to tree felling;
marking
of fruit trees;
ensuring directional
felling of target trees;
the use of aerial log extraction, rather than

skidding, to reduce
chosen
not been
avoidance
the
damage
post-felling
of
treatments such as the cutting
company and may reduce
by the logging
from being able
wood
Gabon.
clear, as there
evidence both
is
of
in
various parts of Borneo and Sumatra.
where hunting
is
orangutans
in
be determined by the quality

in
are
terms
many
of
fruit
fruit
In
areas
not a serious threat, the survival of
productivity.
trees,
and does burn;
fire
appears
of the residual
to
stand
Even where there
however, logged forest can
has a devastating impact on
orangutans. Chimpanzees are also capable

adaptation to ecological conditions
of great
logged forest,
in
but there are few data on the response of bonobos

to logging
on
its
own. Their greater consumption
of
herbaceous vegetation suggests that they might be

reasonably successful
gorillas.
It
is
In
disturbed forest. As with
hunting and the fragmentation of forest
demonstrate
in
live
production; the timber industry
bonobo
of
timber
at
therefore, the selective extraction
moderate
intensities
from a forest
need not cause the complete loss
of
great ape
populations. The impact could be reduced
companies took steps such
if
logging
as:
in
reduce the impact
wildlife.
few
companies
to
istry of
roads,
canals
le.g.
in
although
model
is
a joint project in Congo,
Wildlife Conservation Society, the Min-
des Bois,
some
in
their areas
Instance, the
oil
prevent trade
in
the
hunting and trade
of great apes. In forestry
tried to control
of
Influence.
to the lack of
bushmeat hunting
Cameroon,
for
tried to police
and
In
company Esso
bushmeat during the construction
Media exposure
and
pipeline
of the links
during
attitudes.
to
be effective
in
2001-
between logging
commercial bushmeat hunting
leaving patches of unlogged forest of sufficient
taking other steps to prevent forest fires; and
in
con-
National Park,"
have pointed
Cameroon-Chad
been shown
preserve a moist microclimate and
company. This
logging
to control
critics
nesses have also
are completed;
to
independent evaluation or monitoring. Other busi-
Bornean
and bridges after operations
19981 to
Forests and Environment, and Congolaise
Industrielle
2003."''
forests),
on
reduce hunting. One example promo-
ted as a successful
between the
of their activities
have worked directly with logging
initiatives
blocking access to logging areas;
size
adopting a 'code of
in
conduct' las proposed by the Ape Alliance
of
swamp
some companies have
indeed taken an active interest
suppressing hunting on their concessions;
destroying
component
therefore seen as crucial to the future
is
protection of great apes, and
cessions near Nouabale-Ndoki
In principle,
a key
In
timber
range states. A positive engagement with logging
companies
bushmeat, especially
destroy chimpanzee and
is
for
the economic and social development of most
more
to
managed
forests
in
partnership seeks
lil<ely
that
friendly' way.
great ape populations, particularly
Central Africa,
In
consumers
to
ape
a 'great
areas by logging roads and settlements that are
populations.
256
to
produced
Many
serious impacts
and remarkable adaptation associated with logging
IS
profit
however, potential benefits
margins; there are,
in
silvicultural
of climbers.
these measures require greater Investment
All
Logging
have
to trees that
and
for felling;
has
also
changing business
The Inter-African Forest Industries Asso-
ciation (IFIA) represents 14 logging
together held concessions on
UO
companies
000 km'
that
of forest
and employed 20 000 people
committed
its
members
the late 1990s.
in
has
It
prevent
to take action to
bushmeat had
much
of
Kahuzl-Biega National Park was not controlled by
established a working group
the park authorities, but
IFIA.
executive
chief
of
European timber companies, including
This aims to promote better forest protection
and management (including protected areas],

enable investment
to
local forest industries,
in
reduce the impact
to
and
bushmeat trade on
of the
vulnerable species. The group
vs/orking to
is
develop
cooperation between private companies, NGOs, and

the World Bank,
order
in
to foster
integrated use of forest resources
Africa.
It
tropical Africa,
also seeking to promote a 'code of
is
conduct' for forestry companies
in Africa,'^
but
NGO
was occupied by coltan

was also being used to
militia." Coltan
finance various armies
was described as
in
DRC
during the war,'^^ so
'conflict coltan', further
strength-
ening the grounds for a boycott. The aim
campaign
was,
DRC
lower
coltan to the electronics
com-
panies that ultimately were buying
different
the
of
in effect, to
to boycott coltan
the value of
sustainable and
in
Congo Basin and West
with an emphasis on the
miners and
it.
approach was adopted by the DIan
Fossey Gorilla Fund Europe
which brought together

sought Investment
in
In
all
the
system
Durban Process',
the stakeholders and

of environmentally
and socially responsible mining outside protected
observers and even the World Bank have been
areas, that would
frustrated at the slow progress and lack of concrete
to
results.
partnerships between business and conservation
tions
this
allow poor artisanal miners
valuable
International (FFIl, for Instance,
from producer
consumer,
to
like that
have been introduced
recent years. These are
in
promoted by NGOs (such as
WWF,
Greenpeace, and
mechanism
Friends of the Earth) as a
for tackling
logging and developing sustainable forest
illegal
management. The
members
corporate sector
to
increase the area
of certified tropical forest. Certification
FSC-certified
Malaysia, and one
is
and Southeast
slowly, however, in Africa
are two
works with
Tropical Forest Trust
sites
in
There
Asia.
Uganda, one
however,
rate poorly,
Flora
working with
Is
Global Compact, a UN-sponsored network,
promote responsible corporate
citizenship,
to
develop markets for ethically sourced coltan as
an investment for peace
In
DRC." The aim
is
to
persuade large western companies that use coltan

(such as
Motorola, Sony,
Hewlett Packard, and
Nokia) to buy the mineral only from environmentally
and socially responsible operations
In
DRC.
in
Coltan (tantalum ore),

confiscated by
at
Forest restoration
patches
in
of
unconnected
wardens
Kahuzl-Biega
National Park,
among
which they
the Congo.
Great ape populations can be fragmented
Ian
management
addressing wildlife
UN
New
developing
Sumatra.'^' Existing certifica-
in
schemes generally
the
to
are
resource."
now emerging. Fauna and
groups
timber through a chain
developed by the Forest Stewardship Council (FSCl,
tion
still
and schemes
of custody'
its
from
benefit
Environmentally responsible logging operato track
play
impact on gorilla and
a devastating
other wildlife populations.'" Until 2004,
commercial hunting.'" The World Bank has also
officers of
in
habitat, within
Redmond
issues.
Mining
A comparable approach has been adopted by
inter-
national conservation groups to the certification of
mined materials, and

use.
to
For example,
paigns have had
advocacy surrounding their
cam-
international advocacy
some
effect In stopping
companies
from purchasing coltan from Central Africa."
As discussed
In
Chapter
from which tantalum

Is
extracted:
it
is
(a
used
aerospace alloys and
13, coltan is
an ore
heavy metallic element!

In
for
certain
surgical and
making heat-resistant
capacitors for mobile phones and other electronic

products. High prices led to a coltan rush
with
thousands
of
artisanal
in
2000,
miners entering
Kahuzl-Biega National Park and the Okapi Faunal
Reserve
in
eastern DRC. The resulting
demand
for
257
World Atlas
The chimpanzees
corridors.
have survived
In
now
benefit from
likely
to
Bossou (Guinea!
at
an isolated forest patch, and are

a
corridor project to
Bossou with the central Mount Nimba
link
forest
area.'"'
Local communities
The
loss of food sources following forest degra-
dation and clearance has led

to
forage
In
with
conflict
some
great ape groups
farmers.
'"' '^
fruit Is
scarce
Orangutans also often
fruit
crops adjacent to the forest, and
this
is
may be
becoming an Important issue
Sumatra.
and their
raiders,
Impacts on crops are greatest when

the forest.'
into
Chimpanzees are most
commonly documented as crop

In
them
agricultural areas, bringing
raid
killed;
parts of
In
southern Burundi, this Issue has been
In
tackled through a project that employed a small
group
well trained people to protect villagers
of
from raiding chimpanzees; they monitored the

locations of
chimpanzee groups, and drove them
A tree nursery in
are vulnerable to chance factors. Forest restoration
away from
Guinea, part of a
aiming
been Implicated
program
to create a
'Green Corridor'
size
reconnect such patches or increase their
to
has been
promoted
little
The few examples
states.
in
the great ape range
of successful projects
between Bossou forest
include the two FSC-certified forests
and Mount Nimba
both
in
restoration
Uganda,
to finding
carbon-
communities.
order to link two
storage funds. One
National Pari< and aims to increase the area of
wildlife
populations.
chimpanzee
sufficient
at
habitat.'
and wetland restoration are
forest
rate
of forest,'"
clearance for palm
forest
of
oil
plantations by far exceeds the rate of reforestation.
Connecting forest protected areas and other

blocks
habitat
by
establishing
viability
anzee
to
population and habitat
Uganda estimated
some
of
are
that the extinction risk
Uganda's chimpanzee populations
much as
linked
by corridors.
Is
when popu-
55 percent
If
options
to
between people and
payment
resolvable through the
compensation
for loss,
damage caused
quite tolerant of
of
and people can be

by wildlife
if
they
arrangements
and paying compen-
for negotiating
sation for loss, injury, and death have been tested
since the beginning of wildlife conservation

early years
the
of
century;
last
arrangements are Increasingly coming

ted by both the public
Is
reserve
in
required to give up
its
benefits.
If
it
is
to
be expec-
when
right to
any way; an opportunity cost
Imposed that must also be

of
the
and the conservationists.
Matters become more complex
munity
In
benefit-sharing
offset by
com-
use a nature
is
then being
some
other set
not to cause friction. Diverse
maintain large populations are limited, ensuring the
options are available for achieving
opportunity for individuals to disperse
agreements. These range from easement contracts
Is
critical.
sustainable
Highly fragmented populations of great apes that
to
might benefit from the creation of corridors Include
logging, or farming) while allowing for continued
the Cross River gorilla

of
in Nigeria,^'
Sumatran orangutans
Indonesia."^
On
In
and populations
the Leuser Ecosystem,
broader spatial scale, the West
African chimpanzee population would benefit from

habitat restoration
258
impacts on local
to limit the park's
conflicts of Interest
are
Uganda,"
In
significant attention
stand to benefit from wildlife revenues. Various
assessment (PHVAI on the eastern chimp-
in
reduced by as
lations
and protecting
has been proposed by a number
'corridors' of forest
of conservationists^'' '"
Most
Kibale
Kinabatangan, Sabah,
In
underway, countering the ongoing loss

but the
ways
in
located within the
and crops." Gorillas have also
crop raiding, for example outside
Bwindi Impenetrable National Park
projects financed with

is
in
where managers have devoted
chimpanzee
efforts
villages
and the protection
of existing
compensate
use
of the forest for collecting
other items of
to
outright
often
little
land
however, which
is
for certain lost rights (e.g. hunting,
Is
medicinal plants and
conservation concern, through
purchase. All require funding,
why
international donor support
so welcome, and
why
benefit-sharing
schemes
in
play
(based, for example, on gorilla tourism)
are so attractive as a self-financing alternative to
grant
Nevertheless,
aid.
who
that people
is
it
important to recognize
give up land rights so that the rest
can have a nature reserve have earned
of society
the right to
be paid
fairly.
Participation
in
the
protection of a natural landscape that produces
clean water, flood control, soil protection, carbon
sequestration, and
great ape
deserves
benefits
appropriate remuneration.
INTEGRATING APPROACHES
has increasingly been recognized that biodiversity
It
conservation and sustainable development are

inextricably linked. Consequently, addressing the
most immediate concerns
of
local people
living
around protected areas, such as improving health

care and access to education, has become a major
'" '"
Two common
feature of conservation policy'
approaches that attempt
to link the
conservation
Iroko Foundation
of natural
of
local
resources and the development needs
people are Integrated conservation and
Korup National Park and Dja Faunal
the
development projects (ICDPsI and community-
Reserve
based conservation. These both aim
the long-running Lake Tanganyika Catchment
erty
to alleviate
pov-
and make rural livelihoods more sustainable,

""'-^^'
in and near protected
especially
promote
typically
activities
such
Cameroon;
in
west-
ern United Republic of Tanzania (and others
as:
founded by the same
Mengame Reserve
growing trees along with
i.e.
institution,
project
Nyalama
the project at
In
in
such as the
Cameroon];
Guinea", which
Is
other crops for building poles, firewood, char-
funded by the European Union and the US
coal, silage, etc.;
Agency
tree nurseries, for species bearing fruit
other foodstuffs,
fire
wood, and palm
in
the Niger Delta.
Reforestation and Education (TACAREl project
beekeeping;
agroforestry,
African giant snails for

sale on the roadside
founded by the Jane Goodall Institutes
areas.'-
Such projects
in
for International
Development; and
the Leuser Development
and
Programme
In
nor-
thern Sumatra.
oil;
sustainable farming, for cash and subsistence
an ICDP
between
crops;
At
ecotourism, where appropriate;
various stakeholder groups, with the aim of meeting
environmental education;
all their
its
heart,
is
a transaction
needs, typically including those
of:
family planning;
clean water;
official
low-fuel stoves;
participating international
health care; and
biodiversity;
microcredit loans.
local
donor agencies
to relieve poverty;
NGOs
conserve
to
NGOs and community groups
to
achieve
greater self-sufficiency, autonomy, and colIf
hunting
Is
allowed,
it
is
typically
managed through
lective wellbeing;
and
governments
licensing procedures, harvest quotas, monitoring of
national
animal off-takes, and enforcement. ICDPs with
and international commitments.
to deliver
on
political
objectives that Include the protection of great ape
populations Include those
for:
Fulfilling
all
these agendas
harmonious way
the
Oban and Okwangwo
River National Park
Divisions of Cross
in Nigerla;^'''"-^^
in
view
Is
extremely
In
balanced and
difficult,
of the different scales of
particularly
time and space by
which each group judges success. Donors want
to
259
World Atlas
prowess and dietary
and expense, with a favorable mid-term and post-
groups, through certain taboos. The fact that a wild
project
National governments want
evaluation.
significant benefits delivered quickly to electorates.
Local people want
and
hopeful
to feel
about
NGOs
Conservation
more secure, empowered,

long-term
their
future.
want a protected area
often
with clear and stable boundaries, that
is
safe from
diversity and, for
animal can be trapped

appreciated
may be
wild-caught meat
market
at prices
meat.^'
The 'bushmeat
available
the fact that

in
the local
West and Central
crisis' in
result of these preferences
by a growing
being expressed
improved transport infrastructure and access
Some
staff.
variations
rare
ICDP
that
it
is
everyone. The most
satisfies
remote areas
full of wildlife.
population with
This trade
is
to
serviced
by energetic hunting and trading fraternities; like

specialists in any other
commodity, these are often
based and possess secretive commercial
successful tend to be those that are slow acting,
clan
forum based, and that create genuine partnerships
contacts over large areas. To the extent that the
of
common
interest
NGOs,
international
and understanding between

local
NGOs, and community
groups - with the agendas
both
of
external
government and donor agencies sometimes being

given secondary importance.
In
other words, to save
ecosystem and the great apes within
a local
it,
local
bushmeat trade
these groups
ongoing, and cumulative
more
fully in
Chapter
in effect,
as
is
discussed
People
in
who
live in
Africa] are
used
to
consuming meat
from many vertebrate species. They may define

their
identity
partly
in
terms
of
as criminal gangs
and law enforcement mean that not

hunters realize that their
all
great ape
activities are illegal.
Early attempts to curb this trade
on the
said,
were founded
people consume bushmeat simply
belief that
protein,
and
sell
bushmeat
just
the context of integrated
In
conservation and development projects IICDPs), the
CRISIS
hunting cultures (such as those
the non-Muslim areas of Borneo, West Africa,
and Central
is
and shady business. That
because they need cash.
MANAGING THE BUSHMEAT
for great apesi,
it
of
both rural conservation education
in
because they need
15.
las
is illegal
may be thought
that run a lucrative
weaknesses
transactions are needed; these must be continuous,
their
hunting
Jobogo Minndi/Virunga
Virunga
also often
is
is
both hunting and harvesting, and from foreseeable
occur around these hoped-for outcomes, but
gorilla in the
free
lower than are charged for farmed
emerges as
Africa
for
a rural setting, as
in
ethnic
threats that cannot be resisted by well trained and
well equipped enforcement
Rangers and a poached
some
design and implement a project of limited duration
NP
idea arose that the provision of alternative sources

of
both protein and cash would allow effective en-
forcement programs
in
the prevention of this trade.
Protein projects' have therefore been developed to
provide a nutritional supplement or an alternative to
hunted
wildlife.
with animals to raise for meat
le.g. rabbits, cattle,
or
chickens!, or support for keeping quick-breeding
National Park,
and culturally acceptable species. Examples
these include cane rats [Thryonomys swinderianus]
the Congo.
of
and African giant snails [Archachatina marginata].
The Developpement d'Alternatives au Braconnage

en Afrique Centrale IDABAC)
project
(known
in
English as Development of Poaching Alternatives

in
Central Africa) operates
states of Gabon,
becoming
in
the great ape range
Cameroon, and Congo.'""
It
is
now
clearer, however, that a regional bush-
meat issue cannot only be addressed

and must vigorously
sources
of
hunting bans
in
such ways,
target:
supply through, for

in
instance,
logging concessions and the
closure of disused access roads;
market demand through information and education
and
260
Rural families have been provided
campaigns delivered through schools
all
media,
to
replace the concept of
bushmeat as virtuous with

dirty, primitive,
trading
that of
and frequently
and
through the treatment
links
bushmeat
traffickers of illegal
H.3
Table
Gorilla tourism revenues in East
and Central Africa"
the
of
any other
like
Period
Site
Visitors/year
Volcanoes National Park. Rwanda
bushmeat
parts of Ghana, the consumption of
awareness campaign.'"
Kahuzi-Biega National Park,
Akwa
mans
'dirty
1985-1989
800
525 000
1986-1990
800
250 000
1988-1991
2 000
200 000
1994-1996
2 800
450 000
1995-1996
1200
60 000
Uganda
Ibom, occupational hunting
and poaching has become a
DRC
Bwindi Impenetrable National Park,
the Nigerian states of
In
DRC
declined dramatically after an intense and pervasive
Cross River and
Revenue/
year 1US$)
criminal gang.
In
play
being
it
illegal;
in
Mgahinga
Uganda
job',
Gorilla National Park,
leading to a drastic decrease, over the last decade,

in
the
number
reduction
These exam-
of full-time hunters.^'
ples offer important clues to
how
permanent
hunting pressure might be achieved,
in
through an integrated process that brings a whole
range
focus on achieving cultural
of influences to
change, including explanation and outreach, prohibition
and enforcement, community mobilization,
In
some
countries, such as Uganda, gorilla-
based tourism
is
seen as a means
of alleviating
poverty and as an ideal conservation tool;

to
attract
sufficient
visitors
"' In 2004, both
investment.'"
it
is
able
merit significant
to
Uganda and Rwanda
and investment for sustainable livelihoods.
increased the price of an individual gorilla-tracking
GREAT APE TOURISM
with a family of mountain gorillas, making
permit from US$250
Where nature-oriented tourism contributes

way
sustainable, benign
local people,
but there
is
wildlife,
Numerous operations
great ape tourism claim to
meet
benefit. Great
it
known
is
that provide
this strict definition,
often a question about
communities
support the wellbeing of
to
ecosystems, and
as ecotourism.
in
how much
local
ape tourism generates
most expensive
US$350
to
authorities,
demands
and
is
is
it
the
in
the
usually subject to
of national
and subnational
also vulnerable to changes
in
Uganda, the distribution
of
conservation costs.
income from
one hour spent
wildlife-viewing experience
world.'" The income, however,

the conflicting
for
In
tourism
gorilla
at
Mgahinga
Gorilla
National Park has varied significantly according to
and tracking fees;
national policy. The proportion distributed to local
during 1985-1998, the annual income from gorilla
communities has been consistently small;^ there
income from entry
tourism
Revenues
some
is
ranged from
has been conflict both within the Uganda Wildlife
000' '" (see Table 14.31.
Authority lUWAl over the distribution of ecotourism
individual protected areas
in
US$60 000
are
fees, permits,
to
US$500
over
parks from gorilla tourism
to individual
of the
highest
in
the world, ^^ and
sustained despite price increases. Great ape
tourism
is
well developed for mountain gorillas
three range states; for chimpanzees,

in
It
demand
it
is
Tanzania, Uganda, and (before the war]

is
also
confiscated orangutans
in
Malaysia
channeled
into
considered
to
encourage community support
and that
will
for conserving great
The Convention on Biological
Diversity has
produced guidelines and case studies

the
sustainable
tourism activities
While ecotourism
ties
may
benefit local
communi-
by bringing improved road access or employ-
ment
opportunities, these are often not recognized
to
promote
and management
tial
for
local
ape-based tourism
to
secure benefits for
communities can be further limited by factors
such as war and unrest that cause tourism income

to fluctuate.^'"
Nevertheless, hundreds
Bwindi Impenetrable
of
National
people
Park
live off
in
the
Uganda,
of
where
foreign tourists trek to view gorillas and
vulnerable ecosystems and
where
local people
planning
in
UWA."
communities imposed by conservation.' The poten-
apes as well as other charismatic species and their

habitats.
the
a
to
be one poten-
in
generate revenue that can
conservation,
budget allocation
as compensating for the perceived costs to the local
to
is
way
sometimes viewed the income from tourism as

justification for reducing the
many
other range states

sustainable
and
Ugandan government. Central government has
DRC.
partially
Tourism
be
in
of
for
land, increasingly, in Indonesia).
tially
available
the viewing
provided
rehabilitated,
in all
UWA
income among parks, and between the

the
habitats of major importance for biological diversity.
camping
staff,
work as rangers, guides, and
or sell food, crafts, and entertain-
261
World Atlas
also those near
from
far less
Ketambe have generally
suffered
than have other nearby
illegal logging
areas. This effect presumably arises thanks to both
where
the greater presence of outside observers
the tourism occurs, and the benefits of tourism that
might be valued by those who would otherwise be

logging or hunting
in
the area.
Tourism cannot protect
all
populations of great
shown
apes. Analyses of tourist revenues have

it
highly unlikely that the costs of
is
protected areas
generated
in
in
that
managing
Congo Basin can ever be
the
way.'" Mountain gorillas
fully in this
live
small habitat islands that can be both easily
accessed by tourists and intensively protected by

small numbers
relatively
They may be
of rangers.
able to survive with the support of ecotourism, but

this
mechanism
is
unlikely to support the conser-
vation of eastern and western lowland gorillas
and
Stuart Ctiape
other great apes that occur over wide ranges and

Ecotourists traveling by
ment
boat through the
Valley just outside Bwindi,
Kinabatangan area
in
Box
to the tourists (see
8.i]. In
many new
nesses now offer goods and services
Other ecotourism
Malaysia.
Buhoma
the
local busi-
Africa
in
as part of a range of
If
in
carefully developed
activities,
however,
could
it
contribute toward the costs of conservation and
to visitors.
opportunities
difficult-to-access areas."'"
include the chance to view:
bring infrastructure development that could benefit
both conservation work and local communities.

Tourist spending generally contributes to the

National Park
Park
in
Dzanga-Ndoki
in
CAR"'"' and Lope National
in
in
Kibali National
Park
in
Uganda;"^ and
in
and tour operators, as well as on imported
Tourism
to
famous mountain
National Park
on investments by international hotel chains,
airlines,
goods and services used
Gabon;*'
chinnpanzee tourism
the
profits
gorillas
in
the Volcanoes
Rwanda.""
is
areas
in
the tourism industry.
a fickle business;
few ecotourists travel
at risk
from war or
19V0s
the African Great Lakes region, however,
in
demonstrated that
In
Southeast Asia, ecotourism has grown up around
several orangutan rehabilitation centers
and Indonesia, but lack
of controls
in
Malaysia
and poor plan-
ning has led to serious criticism of the practice'^''

(see
Box
Central
Tanjung Puting National Park
14.11.
Kalimantan and
Sumatra are unusual
in
wild orangutans deep
Bohorok
in
research
northern
the forests. At Tanjung
in
Camp
ters at Sungai
Wain and
who
Leakey, the long-term
There are plans
site.
in
that tourists can observe
Puting there are also free-living rehabilitants
spend time around
"'
for
ecotourism cen-
at Kutai National
Park
in
East Kalimantan.
Another way
ask whether
forests,
it
quite
forests around
262
gorilla
tourism
durable. There are always at least
prepared
to
remarkably
enthusiasts
in
the hope of a
gorillas.
The temporary
accept certain risks
meaningful encounter with
is
some
Rwanda
collapse of the gorilla-watching tours to
and
DRC
during the 1990s due to the
genocide there led

gorillas in
gorilla
to
civil
wars and
an increased demand
to
view
Uganda; as soon as the fighting stopped,
tourism began
to
pick up again
Rwanda and DRC. The world now

wealthy people who are prepared
to
both
in
many
contains
pay
to
en-
counter semi-wild great apes; they will keep

coming, as long as the apes are there.
Even well planned ecotourism has potential

to look at great
ape tourism
has actually contributed

apart from
its
answer
to this
Bohorok
in
may
is to
to saving
benefits to
administrations and local communities.

sible that the
or where
civil strife,
there are severe health problems. The turbulent
It
is
costs, however, in particular the risk of disease
transmission between
park
Chapter
pos-
merely another form
be yes, as the
northern Sumatra and
8|.
humans and
apes"'
''
(see
Great ape tourism can also be seen as

of exploitation for entertain-
ment and commercial

debate over whether
gain; there
it
is
is
an ongoing
morally acceptable to
subject any wild ape to
ape
to potential or actual
Table 14.4 Captive-breeding programs for great apes
harm."
Taxon
out
is
principles of zoo-based conservation are set

in
AZA,
spp.)
Western chimpanzee [Pan troglodytes verus]
Overview
Studbook
Authority"
Chimpanzee [Pan troglodytes
CONSERVATION EX SITU
The
ARAZPA
regional
EAZA
regional
Bonobo [Pan paniscus]
EAZA, AZA
international,
Western
EAZA, AZA,
international,
regional
the World Zoo Conservation Strategy,^'' which

gorilla [Gorilla gorilla gorilla]
currently being revised. Publication of the World
Zoo and Aquarium Conservation

pected
in
2005. Both
Strategy'''' is ex-
documents emphasize:
ARAZPA
regional
Eastern lowland gorilla [Gorilla beringei graueri
Bomean orangutan [Pongo pygmaeus]
international'
EAZA, AZA,
international,
ARAZPA
the need for zoos to maintain populations of
animals
(to
minimize loss
and maximize retention
regional
ARAZPA
international
ARAZPA, Australasian Regional Association

EAZA. European Association
educational rote of
of Zoological
Parks and Aquana;
AZA, American Zoo and Aquarium Association;
of natural behaviors;
the ambassadorial and
EAZA, AZA,
abelii]
of genetic diversity
of natural behaviors) in
conditions that promote good animal welfare
and allow expression
play
any practice (including
habituation and tourism) that intrudes and exposes

that
in
Only a small
program
number
for tfiis
of
Zoos and Aquana.
of old eastern lowland gorillas are present
subspecies
is
in
zoos, and an active
not possible.
such populations;
that populations of captive animals are val-
uable
in
furthering knowledge of the biology of
genetic factors such as avoidance of inbreeding or
species; and
founder contribution
that such populations should link with con-
factors such as the age or social history of the
the wild, whether through
animals concerned. The species coordinator also
servation activity
in
to the population,
animal-management issues
and social
breeding for reintroduction or raising funds
identifies
for conservation in situ, either by the holding
be addressed or researched. These might include
establishment or by other organizations
investigation
more important than the former

species held
in
(it
is
is
much
for
most
increasingly clear that the latter role
zoos).
into
causes
of
that
death or
need
illness,
to
or
research into reproduction or social biology

David W. Liggett IvLwwdaveliggettcomI
Captive great ape
breeding programs are
Management
of captive populations
The management
apes
is
of captive
taking place
populations
in
many
great
zoos, including at the
carried out through regional collaborative
Columbus Zoo and
breeding programs
managed by
of
the American Zoo
Aquarium, v/hich
and Aquarium Association (AZAI, the European
houses the bonobo
Association of Zoos and Aquaria (EAZA), or the
Australasian
Regional Association
Zoological
of
pictured here.
born
She was
in
2000.
Parks and Aquaria (ARAZPA). All use studbooks as

their basic
All
management
species
tool (see Table
other than
managed as separate species

AZA manages chimpanzees of
A.4).
chimpanzees are
or subspecies.
all
The
four subspecies
EAZA (which has a

number of Pan troglodytes verus in
its region), manages that subspecies as a separate
population from chimpanzees of unknown origin or
mixed subspecific status. Movements of animals
as one population, while the
relatively large
between
different collections typically take place
on the recommendation of the designated species

coordinator,
tors
in
who takes account
of a
number
of fac-
making recommendations. These include
263
World Atlas
Box U.I
ORANGUTAN TOURISM
Most experts therefore recommended change.
Some argued
Orangutan tourism has operated continuously
since the early 1970s.
From
the outset,
on rehabilitant orangutans,
locused
it
orangutans cap-
i.e.
managed
educational benefits."'
and had been,
reap both economic and
Others argued that
visitor
and the benefits promised were rarely
control
and any benefits simply
tured for the illegal wildlife trade as infants, then
realized,
confiscated from illegal captivity and rehabilitated
the costs to the readaptation and health of the
to forest
Four rehabilitation sites have been
life.
heavily involved: Sepilok ISabah), Tanjung
Puting
Efforts
sites,
had conservation
local
this
source
generate benefits
"' This
to both.'^'
because
it
proved highly
it
is
far easier
view rehabilitants than wild orangutans. Wild
to
orangutans are semisolitary, elusive, and

stay high
typically
the forest canopy; rehabilitants are
in
habituated
to
ground, and
visit
accessible feeding sites
daily,
and
and rehabilitant-
rehabilitation projects
focused tourism were launched,
orangutan readaptation was

little
thought was given
tourism. These
of
to
became
began assessing orangutan

1970s.
late
Two
sites
knowledge
relatively limited
of
and
the negative impacts

evident
when experts
rehabilitation
the
in
were already experiencing
heavy tourist usage by then; Bohorok attracted up

5
to
000 visitors annually; Sepilok drew up
to
undermining
of feralization of the ex-captives.^''^
Perhaps more seriously, tourists and

habilitation projects
were shown
to
staff at re-
be sources
of
diseases (such as tuberculosis, hepatitis B, and

poliomyelitisl that could pass to rehabilitants,
then into the wild population.^'
and
'""
revenue."
other words, the reha-
In
encouraged
rehabilitants being
display rather than to
based
that
captives.
to stay
it
the problems identified
of
into the
because the
990s and
initial
the turn of the century, a disturbing
it.
These media include
static infor-
mation boards, interactive computer-based presentations,
short films, and oral presentations given
by keepers, volunteers, or education personnel.

2001
EAZA
In
ran a year-long campaign on the issue of
bushmeat, with the aim
of raising
awareness and
funds, and organizing a petition to African leaders
for
picture of
orangutan-focused tourism was emerging, with

little
evidence to support claims of economic or
educational benefits, and growing worries about
adverse conseguences
One
of
to
few
the
both health and behavior
systematic
on
studies
orangutan tourism was a case study
at
Tanjung
Puting focused on tourist-orangutan interactions

'"
rationale.
The expecta-
and the educational

tions
and
behavior
of
the
tourists
were not
tempered by the minimal educational programming provided. Many
of
much
them
like
interpreted young
human
control of the illegal
the reasons for
the
recommendations
and the European Parliament
media conveys
in
some worsened,
change 'rom experts had not been adopted. '^^ By
ex-captives to be
the endangered status of apes and
sites
greatly intensified the costs to ex-
Many
Great apes are a focus of attention for zoo visitors.

Interpretation through a variety of
for
forest-
such an uncontrolled
in
Education and the 'ambassadorial' role
messages on
around
orangutan-rehabilitation
at
970s persisted
often
be driven by
interests, with
resume independent
soared after the early 1980s
to
life.
Tourism
17 000.'^ Tourism-related problems that surfaced
included excessive rehabilitant-human contact and
of
orangutans was coming
economic rather than conservation
manner
on schedule.
When
bilitation of
humans, comfortable near the
some
discontinue tourism at
to
'"^
businesses had become dependent on
and
the expectation that
change
for
tourist-rehabilitant contact. '''
were made
would
in
not offset
but this proved difficult because the sites
education and fundraising as secondary aims, and
promoted tourism
did
One recommendation
restrict
to
[Central Kalimantan), Bohorok (northern Sumatra],

ISaravi/akl. All
'"'
orangutans."'
was
and Semenggoh
attractive to tourists, largely
26^
that tourism could be,
effectively to
bushmeat
to
infants
demand
trade.
and
better
The campaign
information boards were seen by millions of people
across Europe;
it
raised
about US$50 000 and
collected 1.9 million signatures on a petition to the
European Parliament. The

in
a report on the
latter resulted directly
bushmeat trade being adopted
by the European Parliament

report recognized the
in
January 200^. This
bushmeat trade generally
as a livelihood- and poverty-related issue.
recognized that
illegal
trade
is
major threat
It
also
to the
sought every chance
hold and cuddle them,
to
unaware or unwilling
either
acknowledge the
to
were not
orangutans,
potentially
wilder areas.
pressure on
increasing
later survey of other rehabilitation
confirmed that tourists were rarely informed
sites
of
and sought out wild
'real'
issues
rehabilitation
or of
how
they should
We know
no systematic studies on the
of
economic impacts
of
orangutan-focused tourism
There has been, as
large
by
reach
benefits
local
few
businesses;
(external]
communities.
very
systematic
little
of
tourism on
the success of ex-captive orangutan rehabilitation;
disease transmission from tourists to ex-captives
and from ex-captlves

of
to wild
orangutans; eco-
orangutan tourism on local
communities; and educational impacts on both the

tourists
and the
Anne
E.
local people.
Russon and Constance
L.
Russell
of wildlife
tourism elsewhere: most economic benefits are
captured
yet,
would be most welcome on impacts
on local communities. Available information suggests patterns similar to those typical
acceptable, but
research on orangutan-focused tourism. Research
nomic impacts
behave with ex-captlves.'^*
play
only with very careful controls.
health hazards this created. Others insisted that

rehabilitants
was considered
wild orangutans
in
A Bornean orangutan
Centre
in
at the Sepilok Rehabilitation
Sabah, Malaysia.
International
Elaine
fwlarstiatl
operators may, for example, advertise high-priced
ecotourism experiences
sites (costing e.g.
addition
in
visit,
money from
but
orangutan rehabilitation
for a 12
day
Some
International airfare).
to
associated work reaches local people,
salaries
local
at
US$3 600 per person
and costs
Indonesia
in
and
[vlalaysia are
very low by international standards, so
much
income tends
of the
to
remain outside
or even national hands. Local businesses
local
(e.g..
hotels, restaurants, guides, transport, shops! have
around some
flourished
uncontrolled and poorly
but
sites
managed
in
such an
fashion that they
have exacerbated health and behavioral problems

for the
rehabilitants.
and aggravated the deteri-
oration of their habitats.
'^^
Recognizing the extent of these problems and

the lack of evidence of benefits, the
of
orangutan specialists
no longer be allowed
that tourism
to forest
life,
or
great
measures
to
in
areas with
where the orangutans are
eligible for reintroduction."'
African
community
recommended
orangutans that have been reintro-
rehabilitant
duced
officially
recommended
and
apes,
help address
Tourism focused on
it
that
be supported by the
European Union through the European Commission.
search
into
tuaries and
for both
conservation status; support of sancrelntroduction projects; and support
rangers and the
management
of
protected
areas. The involvement of zoos varies too;
Links to conservation
An Increasing number
servation
states.
In
in
raise funds for particular projects
in situ
of
zoos contribute
the field by supporting projects
The support provided may be
kind,
to
In
con-
range
financial, gifts
visitors,
funds.
in
ecotourism
so supported Include
projects;
research Into the bushmeat trade; re-
pilot
while others provide grants from central
A growing number become
actively Involved
the supported projects.
Organizations such as the Zoological Society
or the provision of technical expertise.
Projects
some
from their own
of
London. Wildlife Conservation Society. Antwerp
Zoological Society, and Frankfurt Zoological Society
265
World Atlas
15
^\
()
'^^^^^^^^^^^^^^
|J^^H
Hi^î
-^
^^^^H
''"
î^^^^^î
^^^^^^^^^H
^^^K^^^:jfl
WMmJ
K,>^_y^
^^^^^L^^l
lit
Orphaned chimpanzees
have been involved
confiscated by
years. Others, such as Bristol
authorities and housed
more
in a
makeshift sanctuary,
Lwiro, Democratic
Republic of the Congo.
In field
become
recently
The
projects.
pilot
conservation for
many
Zoo Gardens, have
involved
directly
in
field
EAZA conservation database
contains data from around 20 percent of
[vtfhich
EAZA
c
1
zoosi
shows
that at least 22 zoos support
great ape field conservation projects directly. At
to
over
1999 and the end

Finally,
Is
So
US$530 000
In
dilemma
of
what
do with great
to
apes that have been confiscated or discarded as

pets, performers, or research subjects. In addition,
they can provide valuable opportunities for conservation education
awareness
and
The number
growth
in
of great
apes held
In
sanctuaries
and the
destruction,
habitat
crisis In Africa,
and forest
ape populations.
recent years,"' probably reflecting
logging,
in
bushmeat
concern and
for raising public
for the plight of wild
has Increased
as well as deforestation
Indonesia and Malaysia. These
fires in
which there are now over 50 world-
Institutions, of
wide, hold animals that are either donated to
them
or are confiscated by the local wildlife authorities.
Sanctuaries holding apes are, for the most part,
range states or
in
in
states close to the natural range,
but there are a small
number
outside of Africa,
Indonesia, and Malaysia that hold former laboratory
animals or animals that were confiscated

that have not yet
Sanctuaries
in
In
been repatriated
In
Europe
range states.
to
range states have a number
of roles
great ape conservation:
value between
caring for individual animals, they
in
zoos have successfully reintroduced
attempt
to
do so
for great
providing
by
fulfill
animal welfare;
role in
place
where
confiscated
animals can be held, they help the authorities
implement robust confiscation
apes
the project Initiated by the late John Asplnall,
policies that
are an important part of the enforcement of
in
which Kent-born gorillas from Howletts Zoo were
wildlife
sent to Gabon to join a rehabilitation program for
by providing animals for reintroduction, they
orphans
of the
bushmeat
may
trade. In this case, both
individuals died but a second attempt
is
under-
law and the protection
of great apes;
be able to help restore wild populations;
and
way."' '^'Captive-bred great apes are likely to have
by exposing local people and visitors to great
particular difficulty
apes, they can have an important educational
In
the wild before they are able
to assimilate culturally
transmitted skills such as
Impact, helping people to appreciate the value
foraging, parenting, and interacting with other ape
groups. There has been
more success with
species concerned and
of the
to
understand
the conservation problems affecting their wild
wild-
born but orphaned apes, as described below.
counterparts.
SANCTUARIES. REHABILITATION, AND RELEASE
Animals
Ovenîew
having frequently experienced inadequate housing,
sanctuaries often arrive
in
In
poor health,
Sanctuaries have arisen largely on an ad hoc basis
inappropriate social groups, or solitary confine-
as a crisis-management measure and are seen by
ment, and a poor
some as
five
being of limited conservation value.
against keeping or trading
can only be enforced
has a suitable
In
266
become an accepted
sanctuaries have therefore
of 2002.
far the only
general, and
In
particular Government-approved
in
solution to the
captive-bred Individuals of various species into the

wild.
apes
from that support, which
least 18 projects benefit
amounted
unattractive for endangered species

for great
if
facility in
live
Laws
animals, however,
the confiscating authority
which
to
place the animals
question. The alternative of euthanasia
is
deeply
care.'^'
that
diet.
Indeed, at least four out of
orphans die before reaching expert
gorilla
This presents challenges
do not usually arise
results
In
welfare
at
for
In
management
zoo animals, and
strong focus on animal health ani
sanctuaries.
Some
sanctuaries have been instrumental
the
of
Mountain Wildlife Sanctuary
Afi
Nigeria. This
state,
the
NGO
play
in
an example being
areas,
protected
estabtishing
in
PandriUus and the
Cross River
in
was prompted
by the efforts
Drill Rehabilitation
and Breeding Centre." At least one sanctuary, the
Tchimpounga Sanctuary
Congo,
in
is
developing a
long-term program centered on community-based

conservation
pensary
including
principles,
health
dis-
serve local communities.'^ Financing and
to
long-term planning for conservation of captive

populations at sanctuaries have been
achieve and there
is
need
on the establishment
great ape sanctuaries.
of
These should address
to
difficult
general guidelines
for
with host govern-
liaison
ments, local communities and authorities,
site
location,
long-term sustainability, management
practices,
animal management, and health issues.'"
Rondang Siregar
vided they can be kept alive long enough to be sold.
The number
African sanctuaries
The
great ape sanctuary
first
lished in 1969
in
the
in
Gambia by
Africa
was estab-
Stella Brewer.^^The
Chimpanzee Rehabilitation Trust Gambia continues
to
of all great
more robust
lation of
63 chimpanzees on three islands
resulted
in
for those
found
River
Gambia National Park." '" Many other primate
throughout sub-Saharan Africa;

a
sanctuaries,
encourages
It
their
in
possession of
sponsors,
illegally
held apes.
in
approach
"
and reintroduction."
Workshop Report lists 20

PASA members, and
these 18 sanctuaries
chimpanzees
(of
in
The apes held
June 2003 included 632
unknown subspecies], 67
(believed to be mostly
in
western lowland
and 27 bonobos. The large proportion
panzees reflects their greater
gorillas
gorillas),
of
chim-
ability to survive
treatment and poor captive conditions;
it
may
ill
also
in
in
in
in
Sumatra,
new orangutans from
at
young orangutans have,
in
the past,
thriving local pet trade, with large

illegally to
in
the range
Taiwan has resulted
in
increasing
moved
sanctuaries for care and,
local
pet trade, which
a side effect of
almost always
bushmeat hunting
(see
entered a
numbers being
Taiwan and Thailand. More
of
emerges as
and
Many
robust application of wildlife laws
numbers
to
to take
releasing captive orangutans (see Box U.ll.
graphical range (or both).
come
ceased
tourist attractions, as well as simply caring for
exported
in
Sumatra.
1995. Several centers are
states and
Confiscated great apes typically
in
Gunung Leuser
officially
indicate higher hunting rates or their greater geo-
sanctuaries as infants or juveniles. They are mainly
Malaysia
the wild); four
Borneo and one
Bohorok, an earlier center

in
in
Peninsular Malaysia,
where orangutans do not occur
National Park
centers
2002.'" There are three centers
Malaysia (including one
Indonesia, with three
sanctuaries, 19 of which are
18 of which hold great apes."
in
were
captive orangutans
and Indonesia
than
effectively
for a conservation
that includes rehabilitation
Indonesia.
Southeast Asian sanctuaries
An estimated 600 formerly
and conservation
its
2003
these
of rehabilitation
each could on
PASA's
2000,
under the care
and lobbies more
of
East Kalimantan,
substantial fines and prison sentences
between the
liaison
organizations,
own
in
Pan African Sanctuary
the
networl<,
Alliance IPASAI.
part
in
sanctuaries have also been established
wildlife
formed
at the
Centre near Balikpapan
Cameroon, where recent court cases have
in
orangutans
policies on the confiscation of illegally
held animals. This policy shift has certainly been
seen
and
some range
Orphaned Bornean
is likely
Wanariset Rehabilitation
and monitor a naturally reproducing poputhe
sanctuaries
in
states adopt
continue to increase as
to protect
in
apes
to
in
rehabilitation
In
litation
illegally
held
captive animals being

if
possible, for
and release.
Indonesia and Malaysia, orangutan rehabicenters have formed a network and held a
exchange experiences and
Box U.2). Young animals that survive an attack by
series of
hunters are often considered potentially more valu-
develop improved practices. At the 2002 Orangutan
able as an item for sale than for consumption, pro-
Conservation and Reintroduction Workshop,
workshops
to
it
was
267
World Atlas
Box U.2 SENDJE,
AN ORPHANED
more money than
CHIMPANZEE
20 blue duikers to
by selling
bushmeat market,
the
this
has become too well
understood by the local hunting population. Before
Thousands
every year
of
tons of busfimeat are
Central Africa, and mucfi of
in
from animals
List of
comes
it
are considered to be Vulnerable,
tfiat
the
oil
boom, great apes were seldom hunted
meat, as the adults are heavy

of
chimpanzees and
Endangered under the
orphans
panzees
Endangered, or
Red
consumed
Critically
to carry.
gorillas are
now
for sale to expatriates.
order
'in
to
for
Whole groups
killed to provide
By buying chim-
save them',
the
expatriate
There are many factors that encourage bushmeat
market has stimulated great ape hunting and the
consumption, including cultural
pet trade.
ing
human
tradition, increas-
populations, and expanding access to
forest areas.
There are also many conseguences,
This
in
the story of an orphaned chimpanzee
IS
Equatorial Guinea,
known as Sendje. She was
including the ecological impacts of eliminating seed
named
dispersers, and the outbreak of diseases such as
which she came. Once orphaned, she eventually
Ebola hemorrhagic fever following contact between
came under
humans and
Equatorial
wild animals. Another effect
growing number
follows
light
is
of
is
the
orphaned great apes. What
after the village closest to the forest
from
the care of a conservation project
Guinea;
Brigid
Barry
in
became her
surrogate mother:
the story of one such ape that also sheds
on the role
of the
community
expatriate
in
Equatorial Guinea.
Equatorial
Guinea
petroleum-fueled
increasing
in
has a quickly growing,
economy
numbers
of
that
has attracted
white expatriates.
Many
of
first
heard about
the city
when
chimpanzees
an unusual
in
the
in
the
sight,
this
baby chimpanzee arriving
saw
the corpses of two adult
bushmeat market.
This
is
not
as dead chimpanzees were seen
market a few times a week. What was
these are willing to buy orphaned chimpanzees,
unusual was that one of the adults was a lactating
(even though this
mother.
is illegal],
'cuteness' or to prevent
local people.
either
because
of their
them from being eaten by
By selling one baby chimpanzee
an expatriate, the average hunter
is
able to
to
make
A local employee of the project then

how a live baby chimp had also arrived in
described
the
market and that a
try to sell
it
to the
taxi driver
had taken
it
off to
expat community.
Sendje with carer. Young chimpanzees need constant ptiysical contact.

Lise Albrechtsen
.>.i:v^&^v..>;:^^^^'
268
Later that
they
sum
of
"sorry for her and thought
felt
perhaps they could return her
to
The
the forest."
prospects for a lone juvenile chimpanzee
in
the
forest are extremely poor; without her mother,
she
would begin
and would be taken by
to starve at once,
scavenger within days. After two days
a predator or
these
new owners contacted
Brigid, claiming that
was
they no longer wanted Sendje as she
When she was
of a handful.
brought
much
too
into Brigid's
was
Sendje was only semiconscious. This
care,
perhaps due
seemed more
her condition
diet over the previous
months
eight or 10
mothers milk
for
old.
up
course
As chimpanzees drink
powdered milk along-
the sheen of
quite active.
must be
first
babies. During
fed about three times
months
18
of
life
man
member
of its
'chimp-sitter' this
and
is
difficult.
to tie one's
becomes tedious when the
activity is
of
Sleeping
shoe laces
performed
to a
the
of
she was a potential buyer
to
the necessary vaccina-
jet,
import papers were processed. Sendje

the journey to
live
among
finally
made
other chimpanzees at
the sanctuary.
Sendje's situation
return to the
If
she remains
50 years
Let's
is
not ideal.
She
will
in
in captivity
good health, she might
and could
hope that the future
vation does not
lie
of
to
live
give birth there.
chimpanzee conser-
behind metal bars. Although
Sendje was saved from the cooking

expatriates
never
she knew before her mother was
life
who bought
pot,
the
her at the disco contributed
the growth of the pet trade for orphaned chimps.
for
anzee baby, offers that were rejected and used as

a
way
to raise
an employee
awareness
community
It
is
of the issue
hoped
among
that, with
the
enough
an
rejections, the hunters will return to thinking that
these endangered apes are too heavy to carry out
Sendje
But what
of
of
of
garden was found with plenty
to climb. At nights, Brigid
one other person took turns
and
at chimp-sitting.
Sendje's future? At two or three
she would become a serious
as an adult chimpanzee
is
much
threat,
stronger than
an adult human. There are currently no animal

in
Cameroon agreed
offered to take her to
company, two daytime chimp-sitters
were employed and
sanctuaries
private
in its
in
were administered, and CITES export and
expatriate
With the financial help
of age,
company
three times whether she would like to buy a chimp-
to leave the
other captured wildlife.
years
Cameroon
oil
house with
was unable
chimpanzee, as she did not want any
of trees for
Chimpanzee Rescue Center
Over the following two months, Bngid was asked
bites. Brigid
oil
discussion with European
ear-piercing screeches
this
American
reintroducing Sendje to the forest. After
take her The
killed.
and
locals to think that
the Mondasi
and American primate experts, the Sanaga-Yong
for
not pleasant. Even having a
shower or bending down
constant accompaniment
of
mother
with a furry creature attached to you through a
sweaty tropical night
in
Bata, Equatorial Guinea.
in
many searches and much
the hu-
its
community For
becomes
hope
human
chimpanzee
does not relinquish physical contact with

or another
are
captivity
in
human
require up to three diaper changes. Unlike
babies, for the
recently killed adult chimpanzee
bushmeat market
tions
Young chimpanzees raised

even more demanding than
the night, they
their
months, Bngid concen-
to 18
bacon sandwiches
she was only about
week she regained
became
her coat and
her
to
supplementary vitamins and
of
calcium. Within a
of
that
trated on giving her a diet of
side
be due
likely to
two days
was estimated
It
Lise Albrechtsen
gash on her crown, which she
to a
might have suffered when her mother was shot, but
and beer
play
a group of drunken Europeans at
nigtit.
the local disco bought Sendje for a large
money because
in
Equatorial Guinea, and there
was no
of the forest
and
that the juveniles are not
capturing. Another thing to hope for
is
worth
that the
national laws protecting endangered species will
eventually
be
enforced.
With
Sendje's sad history and future
become among
chimpanzees
these
life in
changes,
captivity
would
the last such experiences for the
of Equatorial Guinea.
Lise Albrechtsen
and Bngid Barry
269
World Atlas
Rehabilitation and reintroduction to the
therefore
proposed
often
and many sanctuaries work toward

Reintroduction
are
viîld
confiscated
for
apes,
this
goal.
often not possible for a variety of
is
reasons, however, since an animal's capacity to
respond
to rehabilitation efforts
variables, including
duration of
Animals with severe
prior captivity.
its
depends on several
age and the conditions and
its
behavioral problems or deformities, for example
from snare-related
cannot be released
injuries,
safely into the wild."
A field
assistant holds
Reintroduction
rarely
is
task.
who has been
of
tranquiljzed for the
(confiscated apes are usually of
journey to the release
There
site in
Conkouati-Douli
straightforward
an adult chimpanzee
There are often questions over the desirability
mixing apes of different genetic provenance
unknown
origin).
also potential for disease transmission to
is
wild populations, increasing competition for scarce
food resources between reintroduced and native
National Park, Congo.
apes, and the possibility of aggression between
New
them."
now be used
DNA
makeup. Mitochondrial
showed
analysis, for example,
National Park;'"
gorilla
illegally
Thai
at
there
the
to
first
Reintroduction
park.'
arity with
In
If
rehabilitant apes
with great care.
now
2003, she
to
In
Indonesia, the national policy
orangutan populations have been

reintroduction project,
it
new
not
resolved to establish the Orangutan Conservation
Forum
lOCFl. At the 2004 Population
Viability
Assessment Workshop
pledged over US$25 000

as
its
in
funds
in
to
and Habitat
Jakarta,
NGOs
support the OCR,
establishment had foundered through lack of
resources. The forum

advice,
is
to
improve
conservation effectiveness.
individuals
social interactions,
dom
and varying degrees
of free-
(from spacious cages to electrically fenced
enclosures),
they are
still
form
of
captivity.
succumb
As with any
ensure that the

to
the
same
that the
money and
effort
would be
better spent on tackling the factors leading to the
creation of orphan apes
in
the
first
place."
These same concerns also apply

location, a
procedure
in
to
trans-
which animals are captured
wild, given veterinary care during a period
of quarantine,
Although sanctuaries provide food, shelter, care,
lost.
critical to
reintroductions are costly and time consuming,'
some argue
from the
Reintroduction and translocation
will
is
pressures that destroyed the original population. As
intended to provide policy
media awareness, and netvworking
is
allow release at sites only where the local
infant.
Benoit Goossens/HELP International
'J
must therefore be chosen
sites
gave birth to a male
270
is
where
human settlements, their familihumans sometimes leads them to be
Any release
four other individuals),

1996.
available
is
aggressive, to raid crops, and enter villages.''
at
Conkouatl-Douli (with
in
or no hunting."'"'
to
orangutans being
are released near
of
chimpanzees
be released
is little
was planned
2004, testing
in
pointless unless a secure habitat
was one
DRC
from Kahuzi-Biega
identify the origins of over 100
This female
analyze genetic
to
that a female infant confiscated in
was an eastern lowland
held
DNA
genetic research and advancing
techniques can
and then released
in
new
location.^'
It
has been suggested that translocation be used as
way
to
groups
in
consolidate isolated individuals and small
of great
apes
into a single larger population
one area, which would be easier
to protect
and
would have a larger and more diverse gene pool
^M
'
One
potential candidate for
such treatment could be
the Cross River gorilla, which
among
disjunct sites
widely scattered
is
Nigeria and Cameroon^' (see
in
There are so many potential drawbacks
Box
7.11.
this
approach, however, including
to
high cost and
its
the health risks posed to great apes newly
in
contact
many
places.
It
programs
at
five
projects currently have
number
have continued supplying food

while others do not provision,
to
in
of
sites.
Some
released animals,
order to encourage
Minimum criteria for both behavior

have been recommended for individual
independence.
with people and with each other, that such an effort
and health
should only be considered as a
great apes prior to release, and criteria have also
clear that techniques and protocols for
IS
It
last resort.
successful rehabilitation and release
must continue
great apes
of
be developed, and suitable pro-
to
play
has been carried out for orangutans
since the mid-1960s, and
release
in
been proposed
for the suitability of release sites."'
Provided that animals are judged to be suitable for

release, survival rates are good.'""
tected sites should be identified for reintroductions.
number
as the
to
animals
of confiscated
will
continue
grow. Captive breeding of great apes for release
into the wild
is
not a cost-effective conservation
strategy at present. For
it
be
to
effective, not only
must animals breed successfully
many problems of
Many believe
must also be
mam
value of captive
breeding of great apes
not as a tool for conserving
is
wild populations, but as a

for biomedical
and
means to
deemed
in zoos.'"' to
demands
the extent that
apes has been carried out
at a
of
African great
number
of sites;
these involve mostly chimpanzees that have been

released onto islands
in
rivers,
An
ex-
the release by the Frankfurt
is
Zoological Society of 17 wild-born chimpanzees on

the 240
km^ Rubondo
Island
Republic of Tanzania,
in
Lake
Victoria. United
the late 1960s.
in
spent between three and a half months and nine

in captivity in
given
rise
Europe.'" These problem' ani-
left to
own
their
devices, but had
by 2004 to a self-sufficient breeding
population of
more than 40
"""
individuals
living
in
There has been one successful release
of
three groups."
chimpanzees
into
mainland
forest, with
25
of
grams
confirmed to have died, and the fate
being
program
initial
The
for gorillas started in
1994
in
survival rates later improved."'
and
of
chimpanzees
Rehabilitation
in
of six
weaknesses
so
it
in
range states have significant
the areas of education and literacy,"
ness
of
conservation issues must be designed to
take account of constraints within the target societies
and educational systems.

Various techniques have been used, based for
instance on the use of comic books and theater
example
of
the latter
is
An
a play developed by the Wild
Chimpanzee Foundation
to
raise
awareness
in
towns and villages near forests with chimpanzee

Singleton/SOCP
Orangutans released
at
the reintroduction site
adjacent to Bukit Tiga
Puluh National Park
In
Jambi province are

subsequently monitored
sometimes Join
in.
Congo; poor
"
This
in
Gabon
was
in
Congo from 1996.
and release
is
an inherently
attractive prospect, of interest to both the

to politicians,
therefore have an educational component.
the African
equivalent
first
followed by reintroduction of gorillas

1998,
of
All
34
been
4.51.
stakeholders, other measures are likely to
be ineffective. Most great ape conservation pro-
- and
released animals surviving, three having
unknown^^ (see Box
is
great apes. Without awareness and understanding
among
Ian
mals were largely
governance,
arguably the most important tool for saving the
The chimp-
anzees were from West Africa originally and had
years
of
and poverty eradication, conservation education
and that generally
require continued supplementary feeding.

ception, however,
Beside the fundamental reform
investment, trade, national economic development,
so approaches intended to increase public aware-
ethical.^'
and release
Rehabilitation
satisfy
research and for the
scientific
apes' educational value
these uses are
but the
reintroduction
that the
solved.
in captivity,
PUBLIC EDUCATION
media and
remains firmly on the agenda
in
271
World Atlas
Uganda
the
Ngamba
Education Centre and
Wildlife
Island
Chimpanzee Sanctuary, which
play a similar role
in
Uganda;
the Drill Rehabilitation and Breeding Centre
southeastern
NGO
by the
which
Nigeria,
is
Pandrillus and
many thousands
chimpanzees, both
has exposed
Nigerians to
of
at
and
drills
former location
its
in
managed
urban Calabar and more recently
in
the Afi
in
Mountains; and
the Limbe Wildlife Centre, Cameroon, which
is
more than 30 000 people annually

and whose outreach program visited more
visited by
11000 students
100 schools and
than
in
2000.''
The Pan African Sanctuary Alliance (PASAl

Michael Huffman
Two WaTongwe
populations. Performances, given
research assistants,
theater troupe
second generation
8 000 people
in
National Park
in
members
of the Kyoto
Ymako
in
French by the
have reached over
Teatri,
17 communities around the Tai
Cote
d'lvoire.
The play uses actors
University Mahale
accompanied by
research team, and a
behavior, the effects of hunting on chimpanzees,
Tanzanlan PhD
and the connections between animal and human
candidate working
communities.
together
chimpanzees through totems and ancestry, depict-
in
the field.
It
music
live
and
local
the
is
the
kills a
portray chimpanzee
emphasizes human connections
ing a conflict that arises
totem
to
between families whose
chimpanzee and
a hunter
who shoots
chimpanzee. The play can be adapted
customs
message
to
to suit different
to
audiences but conveys
that people should not
kill
and eat
apes, because they are closely related to humans.
Conservation education projects are frequently
an important part
of the
where animals serve as
work
of
ape sanctuaries,^'
a focal point for explaining
develop tools
awareness,
hold
chance
to
Malaysians
see wild animals
in
levels.
They
educators from different
CONSERVATION-ORIENTED RESEARCH
Field research has become intimately
conservation
as
efforts,
witnessed the decline
of the great
projects bring
employment
in
and have become
Research
foreign exchange, provide
for local people,
the area of study.
If
have
apes and the
advocates for conservation. '
strong
linked to
primatologists
destruction of their habitats,
some
and focus attention on
the site concerned
protected, the research project and
is
its staff
already
can be a
source of practical support and knowledge to those

responsible.
is
If it
it
and
and national
develop materials.
Urban
Indonesians,
to bring
sanctuaries together to share experiences and
study or survey
seldom get
Examples
their
not protected, the results of the
may persuade
local authorities that
should be.
of
long-term research programs
natural habitat, but visits to sanctuaries provide
that have
stimulating opportunities, especially for classes of
conservation include those at Karisoke Research
schoolchildren. Examples include:
Center
in
Gombe
the
Chimpanzee
River
Rehabilitation Trust, at the
Gambia National Park, which has helped
eliminate
in
young
the education program of the Lola ya
Bonobo
to
chimpanzees
the
in
Sanctuary close
illegal
trade
Gambia;
to
Kinshasa
has the largest group
272
at both local
workshops
to
improve education and public
to
the purpose and importance of wildlife conservation.

Africans,
also
is
working with the Bushmeat Crisis Task Force
been highly
influential in favor of great
ape
the Volcanoes National Park, Rv/anda,'"
National Park
in
Bossou
Tanzania,'
in
Guinea,'"^ '" the Tai National Park in Cote d'lvoire,'"
Tanjung
Puting
Park
National
Ketambe
Borneo,^' and
in
the
in
Indonesian
Gunung Leuser
National Park, Sumatra.'" These and others are
in
of captive
DRC, which
bonobos
in
described
in
the
relevant species
country profiles of this volume.
chapters and
Many
recent re-
search projects seek to involve local communities
the world and reaches up to 10 000 students
in
each year;
conservation education and ecotourism elements
conservation
activities,
and frequently include
WWF and the
the collaboration between
(e.g.
people of Dzanga-Ndoki National Park
in
Ba'Aka
CAR]."
has been hailed by Conservation International
new model
as a
INITIATIVES
(CI)
Indonesian
level of
society'" Apart from preventing disasters, local
Communities may declare protected areas where
people expect the
they have the legal authority to do so and sufficient
tion with
awareness
sale of organic coffee from the surrounding area
so.
advantages
of the
The Lossi area, some 50
National Park
to
km
be gained by doing
southwest
Odzala
of
Congo, was placed under such
in
community was
protection after the
sensitized by
researchers attracted by the high densities of west-
tourism."
gorilla
the
of
potential
of
gorilla-based
tourism project was developed,
pilot
groups habituated, and monitored from 199A
new park
NGOs such
as
CI,
to
encourage coopera-
which has negotiated the
through the international Starbucks chain. A similar
community-driven
initiative Is
leading the protection
Sungai Wain forest near Balikpapan, East
of the
Kalimantan.'""
ern lowland gorillas found there. The community
became aware
These
conservation
local
the context of a global advance
volvement
In
forest
Initiatives
In
exist
in
local-scale in-
management. Throughout
Latin
America, for example, local governments have
onwards; hopes were high that benefits would be
become
obtained by local communities. The LossI Gorilla
zoning,
Sanctuary was however
charging fines; hundreds of municipalities have their
hemorrhagic fever
population
was
by an outbreak of Ebola
hit
2002,
In
devastated,
in
which the
gorilla
and two habituated
groups were wiped out.'"
in
the early lV90s by the people of the
area to link the
Okwangwo
National Park and the
Cross River
Afl
River Forest Reserve
was
state, Nigeria."" This decision
formed by the educational work
of Pandrlllus
in
inin
the area, by the wish to help protect Cross River

gorillas,
and by the general sense
of
wishing to
participate in the process of conservation
tainable development
and sus-
prompted by the investment
from governmental. International NGO, and donor
agency sources
In
planting trees, fighting fires,
and commissions working on forestry
offices
and the environment." Likewise,

of
in
the Philippines,
the 1991 Local Government Code
has authorized municipal governments

of
up
to
50
to set
km' aside as community
areas
forests to
safeguard water catchment and other environ-
mental services. '" They are Increasingly doing

partnership with local and international
this in
NGOs and
community groups. These decentralized arrangements are

Forest
(see
paralleled
in
Africa through the African
Law Enforcement and Governance process
earlier section
agreements,
this
on Regional
chapter).
activities
and
They give Indigenous
peoples, small farmers, foresters, and local envlron-
Fecal samples, collected
Decentralization
Indonesia following the
in
Suharto regime
of the authoritarian
has often been blamed for an increase
1998
in
illegal
in
logging throughout the country," as local officials
became
In
and around Cross River National
Park."
fall
own
involved
managing parks, granting permits, and
implementation
The Mbe Mountains Community Forest was

established
play
tor conservation cooperation be-
tween the central and regency
COMMUNITY
in
able to award logging concessions without
reference to Jakarta.
In
changes are beginning
one respect, however, the
from wild orangutans

for parasitology
and
DMA analysis,
being
processed
at the
Sumatran Orangutan
Conservation
local
Programme's Ketambe
of their
Research Station,
people are best served by conservation, then they
Indonesia.
to
prove beneficial.
government leaders decide that the Interests
have the authority

that
to
was Impossible
take proactive steps
If
in
way
previously. Thus, the regency
[kabupaten] of Mandailing Natal
under pressure from 30 000
In
North Sumatra,
local people
who had
petitioned the Resident iBupatI), proposed the creation of the

1
080
Batang Gadis National Park, an area
kml The
of
park was endorsed by central gov-
ernment and inaugurated by President Megawati
May 200A.'" This new 'bottom-up'
in
national park
273
World Atlas
also contributed substantial funds
in
port the five World Heritage Sites

that country's debilitating
2001 to sup-
DRC
in
during
war.
civil
Since 1992, the European Commission has
committed some US$50 million
to
Conservation and Rational Use
of
systems
Forest
Eco-
Central Africa (ECOFACI. This has as a
in
priority the
dwelling
the program
involvement
in
activities of forest-
its
ECOFAC
peoples."
engaged
is
directly
with the conservation of the western gorilla and
chimpanzee
example, supporting gorilla-
for
by,
based tourism, biodiversity assessments, and

primate censuses
Odzala National Park
in
and Lope National Park

istration of the
in
Congo
Gabon, and the admin-
in
Ngotto Classified Forest
CAR.
in
Com-
Since the early 1990s, the European
mission has also supported the Leuser Develop-
ment Programme
in
the range of the
Sumatran
orangutan, various projects to encourage reform

staff at Kibale National
mental groups new opportunities
Park, Uganda, fiave
to deter
recently received vital
that tiie next generation of great
equipment for work

the field.
in
unwelcome outside
interests.
projects will take full advantage of

of authority over the
to participate
It
and
expected
is
ape conservation
ttiis
(among the most serious threats
of illegal logging
to
orangutans
redistribution
Finally,
in
Indonesia).
the Global Environment Facility (GEF)
has invested many millions
environment.
dollars
of
the
in
protection of great ape habitat areas, typically
FINANCING GREAT APE CONSERVATION

Over the years, significant amounts of money have
been committed
to great
ape conservation or
to
projects that are likely to benefit great apes directly
NGO community
or indirectly. The
spent several tens
of
governments and
official
matched-funding arrangements with
other multilateral donors and lenders, and with
US$30 000
spend up
range).
to
US$53
of
US$1 0.^6
1970s;
donors have joined
in.
of 2000, for
small grants
lin
The US government plans

million during
the
to
2003-2005 on
conservation and sustainable resource use through

the
Congo Basin Forest Partnership, and
US
partner governments. Of the
US$12
this is
be at least matched by other donors and
likely to
contribution,
per year for three years will be
million
disbursed through the Central African Regional
Program
for the
begun
1995 by the US Agency for International
in
Environment (CARPEI, a program
Development with
environment
in
mandate
nine countries
in
to
assess the
the region.
Meanwhile, the United Nations Foundation
National Park Conservation Project
boundary Biodiversity
Interzone Project
274
With
in
all this
surprising to see
the f>/linkebe-Odzala-Dja
in
money being spent,

how little long-term
great apes has been achieved.
momentum
forests:
of the
the vast
exploitation of tropical
investments
mining, and
structure,
unimaginable diversity
unexpected
effects,
of
actions and actors, side
events
(from
wildfires
to
coups), and cascades of destructive consequences
associated with these processes.

with these
hundreds
hundreds
of
In
billions
of
comparison
dollars and
few tens
of millions of people, a
lions of dollars for forest conservation will
is
2003-2007.'^' The foundation
infra-
in
and the almost
logging;
much
in
less surprising,
human
experience, however, and there
partners
might be
private
has contributed US$3.3 million through the Central
NGO
is
It
it
security for
however, when one considers the scale, pace, and
African World Heritage Forest
matched
Uganda, and
Gabon, Congo, and Cameroon.
if
Initiative,
in
Conservation of Trans-
million to the
be spent very wisely indeed
by
relevant
of
grants include US$^.43 million to the Bwindi
Impenetrable National Park and Mgahinga Gorilla
example, committed about US$1 million each year

support a program
Examples
range-state governments.
GEF
US dollars on
more recently
The US Great Ape Conservation Act
to
in
and
bilateral
has raised and
millions of
great apes since the late
of
the Indonesian timber industry, and the suppression
the
difference.
next
US$25
Wisdom
million
of mil-
have
to
they are to
make
increasing
with
is
spent
the hope that
on
great
ape
much more impact
conservation will have

the last.
It
Toepfer,
the
is
in
this context that in
Executive
than
challenge to
US$25
2003 Klaus
Nations Environment Programme, set a fundraising
play
by declaring that an additional
was
urgently needed
adequately the efforts needed to
United
Director of the
GRASP
million
in
lift
imminent extinction from the great
finance
to
the threat of
apes.'^'
FURTHER READING
Beck. B.B., Stoinski,
T.S.,
T.L.,
Norton,
B.,
Rowan,
Apes and Humans: The Ethics of Coexistence. Smithsonian
A.,
Stevens,
Institution Press,
E.F.,
Arluke,
A.,
eds 120011 Great
Washington, DC.
Brookfield Zoo 12001) The Apes: Challenges for the 21st Century. Conference proceedings. Chicago Zoological Society,
Brookfield,
Brown,
http://vTOm.brookfieldzoo.org/contentO. asp?pagelD=773. Accessed June 13 2005.
Illinois,
D. 119981 Participatory Biodiversity
Conservation: Rethinking the Strategy
in
the
Low
Tourist Potential Areas
of Tropical Africa. Natural Resource Perspectives 33. Overseas Development Institute, http://www.odi.org.uk/
U 2005.
nrp/33,html. Accessed June

Caldecott,
J. 11
996) Designing Conservation Projects: People
and
Biodiversity in
Endangered Tropical Environments.

Cowlishaw,
G.,
Dunbar,
R. 12000)
Primate Conservation Biology. Chicago University Press, Chicago.
The Durban Process: Report of the Meeting. Dian Fossey Gorilla Fund and
Dian Fossey Gorilla Fund 120031
Independent Projects Trust. http://vTOW.durbanprocess.net. Accessed June 16 200^.
ECOFAC
12004)
Conservation
utilisation
et
des
rationnelle
ECOsystemes Forestiers d'Afrigue Centrale.
http://www.ecofac.org/. Accessed July 23 2004.
Hacker,
J.E.,
Cowlishaw,
G.,
Williams, P.H. 11998) Patterns of African primate diversity and their evaluation for the
selection of conservation areas. Biological Conservation 84
Kormos,
Boesch, C, Bakarr,
R.,
M.I.,
(31:
251-262.
Butynski, T.M., eds 12003) West African Chimpanzees: Status Survey
and
Leuser Development Programme
In.d.)
The Leuser Development Programme,
http://vTOW.eu-ldp.co.id/.
Accessed
June 16 2004.
Price, S.V. 12003)
War and
Tropical Forests. Conservation in Areas of Armed Conflict.
Haworth Press, Binghamton,
New York.
Rao, M., van Schaik, C.P. 11997) The behavioral ecology of Sumatran orangutans
Tropical Biodiversity^
Rijksen, H.D. 11982)
How
to
(2):
in
logged and unlogged forest.
173-185.
save the mysterious 'man of the forest"'
de Boer, L.E.M.,
In:
ed.,
The Orang Ulan:
Its
Biology and Conservation. Dr W. Junk Publishers, The Hague, pp. 317-341.

Russell, C.L. (1995)
The
social construction of orangutans: an ecotourist experience. Society
and Animals
3:
151-170.
http://www.psyeta.Org/sa/sa3.2/russell.html. Accessed June 16 2004.

Wallis,
J.,
Lee, D.R. 11999) Primate conservation: the prevention of disease transmission. International Journal of
Primatology 20
(6):
803-826.
Wilkle, D.S., Carpenter, J.F., Zhang, Q.F. (2001)
parks and so
little
The under-financing
of protected
willingness-to-pay Biodiversity and Conservation 10
15):
areas
in
the
Congo Basin: so many
691-709.
ACKNOWLEDGMENTS
Thanks
to Ian
Rutagarama
Redmond
(Ape AUiance/GRASPI for review; to Rebecca
(African Wildlife Foundation) for
Corrigan and John Caldwell (both of
comments on
UNEP-WCMC)
for
Kormos (Conservation
material that
CITES
was merged
internationall
into this chapter;
and Eugene
and
to
Helen
data.
AUTHORS
Nigel Varty,
Sarah

UNEP World Conservation tônitoring Centre
Ferriss,
Bryan Carroll, Bristol Zoo Gardens

Julian Caldecott,
Box
14.1
Anne
E.
UNEP World
Russon, York University, and Constance
Box 14.2 Lise Albrechtsen, University
of Oxford,
L.
Russell,
Lakehead
University,
Canada
and Brigid Barry, Tropical Biology Association
275
World Atlas
Chapter 15
Lessons learned
and the path ahead
Julian Caldecott
n theory at least, a great
ape population could
survive indefinitely within a landscape

of large
patches
of food-rich forest
broad corridors of forest
to
made up
connected by
allow dispersal, inter-
spersed with farmlands and prosperous villages

inhabited by people
snares,
who
who
neither hunt apes nor set
receive good health care, and
who do
not mind their crops being raided. The establishElizabeth
A Williamson
ment
reserves,
and various other forms
ecological
restoration,
of public, private, or
com-
munity investment might create such an ideal

landscape.
such a vision
Putting
into
challenging, however, not least because
effect
is
most great
ape range states face significant socioeconomic

challenges. This can
make
it
difficult for
them
to
undertake the organized, long-term social invest-
Tree planting around
ments
Nyungwe
require.
National Park,
nature
of
enrichment planting, education, compensation,
Rwanda.
that
successful conservation
seems
to
For most of the great ape range states, this

type of investment requires external support. The
cost of this proposed 'help'
and expressed
projects.
come
in
is
often monetarized
the form of programs
The financing
of
made up
of
these projects has be-
the chief priority of an industry comprising
government departments, donor agencies, nongovernmental organizations iNGOsI, and consulting

firms. This industry
enough
to
is
a large one, but not large
counter the overall effects of the
assault on tropical moist forests.
It
is
human
fair to
ask
whether we could do better
A wide range
of techniques
oped by conservationists
great ape survival crisis,
have been devel-
to target
in
aspects of the
the process,
some have
been tested and abandoned. The key lesson learned

is
that conservation
programs need
to
be adapted
to
their circumstances and, generally, have to involve a
long-term commitment
what
in
is
the
of
appropriate resources;
almost always needed
way
in
is
an irreversible
shift
which people perceive and relate
each other and
to their
environment.
to
sustainable
conservation project educates and empowers, en-
courages and enables people

their
276
own
to live better lives (in
terms), causes values to
shift,
and involves
Lessons learned and the path ahead
Following these principles,
effective partnerships.
perhaps we should be examining

application
potential
their
the context of the range state societies
in
with regard to their peoples' history and culture,
hopes and
fears, attitudes to forests
and great apes,
and expectations placed on governrment.
are
While doing so, we should be aware that there

many stakeholder groups involved in any con-
servation enterprise that targets great apes, and
groups have different perspectives and
that these
interests (whether conscious or notl. Broadly, these
stakeholder groups are:
educated middle classes

donate
tical
to
pressure
who
of all countries,
NGOs and
conservation
apply poli-
governments, thereby
to their
directly or indirectly providing the
backbone
of
international financing for conservation;
range state governments, struggling
to
meet
their international obligations, to sustain a de-
Ian
Singleton/SOCP
needs for an
gree of national legitimacy and public support,
and eager
for
much
as
external funding as
possible (on the best terms possible!, whether

in
intact
and suitable environment

and disturbance
free of hunting, disease,
if
they are to breed and raise their young suc-
the form of private investment or official
cessfully and, thereby, to survive
in
Conservation
awareness
at
Medan
International School in
Sumatra, Indonesia.
the wild.
donor assistance;
national conservation and forest-management
agencies and their
bound
to
staff, all
implement
management
more or
less duty-
legislation concerning the
of wildlife, protected areas,
and
resources
forests, while usually also starved of
companies and government agencies
with interests
in
of
in
live
in
where great apes
in
live,
of
communities who
and around the forests

often poor
processes and poorly educated
(in
politan terms), yet typically strong

identity,
perceived values, and
what they want
to
all
of
the
their roles,
rights,
if
necessary,
responsibilities,
ships. In this approach, the great
in
to adjust
and relation-
apes are some-
objectified as 'animals of concern', but this
not to forget their Inherent value as
Is
intelligent
population equivalent
It
in
falls,
achieve
in
of
say, Stuttgart in
to,
particular, to conservation
the great apes
In
Germany.
NGOs and
and represent the Interests
dialogue with the other
human
stakeholders.
monetary
in
terms, isolated from urban decision-making
of
partnerships to clarify and,
scientists to understand
rural areas
or
beings and as our close biological relatives. They
the vicinity of great ape
members
some
are however victims, with a collective worldwide
habitat areas; and

local people,
achieved by
and
oil
the building of roads and
in
other infrastructure
is
stakeholder groups working together
hard-rock or alluvial
agriculture, logging,
gas extraction, mining

minerals, and
human
what
with which to do so;

private
Conservation
WORKING WITH EDUCATION
cosmo-
People
group
change
in
an awareness
who value wildlife

their
own
care enough to justify or to
behavior, or at least to feel guilty
about their negative effects on
wildlife.
successful
process of great ape conservation must validate and
in life.
promote the notion that these animals are suf-
The values and aspirations

stakeholders
may
of
any
of
the above
- but do not always - conflict with
the interests of the final key stakeholder group:
ficiently
valuable to retrain from
will
have a practical effect
among stakeholders
great apes,
explained
in
whose
this
natural history has been
volume, with their specific
killing, eating,
or
abusing them, or from destroying their habitat. This
that
if
it
guides behavior
are able to
influence
events on the ground.
Education can encourage and enable people
to
277
World Atlas
develop the mental tools with which to understand
and communities, but
not only their environments
also their personal potential to achieve adequate
and sustainable
livelihoods.
If
processes, then people are
better able to appreciate them.
If
educational
of
likely to
become much
may
economic,
social,
and technological circumstances are

appreciation
lead people to act
in
right,
this
favor of great
ape survival and against the factors that threaten
it.
toward conservation may be ineffective
Efforts
when
they adopt a single tactic
such
isolation,
in
as offering alternatives to bushmeat without also

enforcing hunting or access bans, or offering
lihood support without also
providing
live-
environ-
mental education and enforcement. Conservation

be more effective
likely to
program
if
it
an educational
in
context, will vary according to ecological
and social
circumstances.
the value and plight
apes are an integral part
of great
and how they are presented
is
delivers a balanced
Providing harvests within the reserve. Not
all
protected areas set aside to preserve great
ape populations need exclude

use.
Extractive'
all
access and
reserves, with
harvesting
regimes organized with
can
people,
local
provide a renewable supply of materials such
medicinal plants, thatching, structural
as
wood, climbing palms, and a range

stuffs, spices, dyes, drinks, etc.
meet household needs
food-
of
These either
directly,
or provide
opportunities for the processing of raw materials
and the sale
of
products such as foods or
handicrafts.
Providing harvests outside the reserve. Even
that includes:
if
a protected area is closed to
may have
human access
advisory materials on laws concerning wildlife
and use,
and protected areas - these should be devel-
harvesting could take place for uses such as
oped with
local people, trialed
those
languages
(to
ensure
appropriate
in
(community
that
a
variety of formats
serve the surrounding areas as legal sources
posters, magazines, etc.)
encourage understanding
change
of
ecology and
the perception of hunting and
in
bushmeat consumption;
programs
that
of
mobile wildlife species that
for
bushmeat, dispersers
plants, crop pollinators,
of
local
own development
accordingly, while
may
economically useful
and pest
control.
Providing local ecological services. Protected

forests can safeguard local
encourage and enable
people to analyze their own environments and

plan their
which
above. Both the protected area and
listed
grounds
radio,
in
the buffer zone will act as breeding and feeding
widely distributed;
in a
a buffer zone
meaning], and
clarity of
educational materials
it
the
ill
communities from
effects of drought, flooding,
erosion,
landslides,
aquifers, wells,
fire,
soil
and the drying out
of
and springs.
actively defending their interests against those
Resisting global warming. Forests sequester
of outsiders;
gaseous carbon
marked boundaries
clearly
for conservation
dioxide. Globally, increasing
atmospheric carbon dioxide levels contribute

climate
areas;
significantly to
explanation and enforcement of hunting and
enhanced greenhouse
access rules;
ging
forums
among
for
dialog
and
resolution
conflict
developing sustainable
in
liveli-
managed expectations
to solve their
that
encourage people
own problems through commu-
nity organization, self reliance,

in
community
to
help meet the cost of forest-based carbon
of activities
such as the restoration
of forest at
Kibale National Park, Uganda. There
hoods;
ness
There are emer-
for the global
storage, which are already forming the basis
stakeholders;
assistance
mechanisms
change through an
effect.
and assertive-
considerable scope for proposals by

nity
may be
commu-
stewardship groups.
Protecting genetic resources.
Many
'"
forest
species contain chemicals which are impor-
seeking outside resources.
tant in the ethnobiological traditions of local
As such
fits
program
is
delivered,
arising from the conservation process will be-
come more
easily
apprehended by
Such benefits can include any or

below.
278
community bene-
The precise mix
all participants.
all of
of priorities in
those
listed
each place.
people, and which might be developed as
com-
mercial products via bioprospecting.
Local
people
may
benefit from these discoveries,
provided that adequate contractual provision

is
made
for access
and benefit sharing on
mutually agreed,
and equitable terms
fair,
These can include clearer
(Including the appropriate recognition of intel-
and better
lectual property rights).
immigration or theft
rights of inheritance,
local ability to resist

of
unwelcome
resources by outsiders.
Providing resources for education and tourism.
Even
small
number
of recreational visits or
make
school parties each year can
a big dif-
ference to a community's economic prosperity,

both
from the sale
directly
goods
of
refreshments, handicrafts! and services
This approach
is
fundamentally 'pro-livelihood'
rather than simply 'anti-bushmeat'. The
model aims
to
(e.g.
elements
the
(e.g.
and communities, and the content
of
clarify
basic
and refine the particular
relationship
between educators
of
community
If
local
communities can
be helped to meet their
guidance, accommodation], and indirectly by
environmental and livelihood discoveries, that have
protein requirements
the profile of the area outside the
most impact on bushmeat consumption and on
through domesticated or
raising
realm
conventional decision makers.
of
Improved knowledge could then
rapidly maturing plant
formulate action guides, tools, and
and animal species, the
forest protection.
Local opportunities. Research, management,
be used
tourism, education, training, and other acti-
priorities for extending the
and around a reserve create many
vities in
different kinds of jobs,
to
other locations
In
process adaptively to
other range states.
need for bushmeat
is
reduced.
from which local people

Gordon Miller/IRF
can benefit as their training and experience

increase.
new
These
activities will also
generate
more
information (making ecosystems
interesting to local people), while encouraging
them
skills
as students at educational
enroll
to
institutions,
thereby helping
and technology
to transfer
new
to local people.
Providing a clean and beautiful environment.
Local people benefit from access
scenery and often a lack

pollution,
which can help provide a
of
all
healthy and relaxed
lifestyle.
Nature reserves
are special places; people living near
stewards
natural
to
crowding and
of
them are
unique resources, and have an
of
important role that
increasingly
is
being
recognized.
Preserving traditional values. Cultures are

distinctive
mixtures
technologies,
live.
preserve those
local ideas
and
often
help to
and ways
of
keeping alive valuable

doing things.
ecosystem close
to
who
a
have a broader range
live
within a
nature reserve
options for
of
is
can use their resources wisely
because they
to
determine
own future, without risking the

their means of survival and prosperity.
their
Improving tenure security.
can be used by local people

rights to
of
A nature reserve can

links,
development than others. This
of
all
where
in
Preserving options. People

viable
forms,
art
the ecosystems
which are rooted

their people
languages,
of traditions,
beliefs,
loss
nature reserve
to
obtain legal
occupy or use nearby areas. This may
mean agreeing
to respect
permanent reserve
boundaries, but secure tenure outside will often

give
more
benefits than are lost by doing so.
279
World Atlas
Box 15.1 GREAT APES, CONSUMERS, AND

THE MEDIA
Over the
tiave
help protect mountain gorillas
to
and have been spent on paying
perhaps one
15
from
25 years, many millions of pounds
last
been raised
Rwanda
This
of the
most famous pictures
sequence
a television
two dissimilar
of
mates together But what has
that
Are mountain gorillas better
off
fame
pri-
achieved''
thanks
David
to
Have the animal stars actually
Attenborough'^
the
benefited from that publicity, from Life on Earth
guards' wages, supplying uniforms and equipment,
onwards? There's no question the world knows
in
and operating educational programs:
come
have
willing to
pay
Government
them.
brought
to
see the
to
more
who are
gorillas, thus
persuading
backing
conserving
to give its full
There
millions
the country from visitors
to
to
no question that the publicity
is
them by
television
has been crucial
in
bringing these things about.
more about
acted
gorillas than
it
did and, arguably, has
response.
in
As
result of this
great apes should,
media
actual conditions are discussed

in
this
attention, all the
theory, be better
in
off-
Their
depth elsewhere
volume, but two issues are particularly worth
considering here.
Many
David Attenborough
now broadcast
television stations
quality wildlife
high-
programs, but these often make no
John Sparks/naturept com
reference to conservation. Yet the eastern gorillas of

been devastated by the mining

found
riverbeds
in
Congo IDRCI have
the rain forest and
in
mineral
of coltan, a
consumed
by mobile phone and computer hardware
compa-
nies servicing the global TV-viewing public. South-
east Asian
palm
we
are also
forests
plantations,
oil
deaths
of
donating
On
from efforts
to
have emerged
improve livelihoods:
If
boost
plywood,
we
are
lin
practice! contributing to the
in
orangutans - perhaps even while also
money
to
television,
area.
project
for conservation
converted to
shampoo, employ
who uses Indonesian
theory and probably
Two key lessons
being
their wildlife destroyed.
eat processed foods, use
builder
WORKING WITH LIVELIHOODS
and
help conserve them.
commercials
for
The development
livelihoods
in
palm
oil
activities
pro-
aim
to
the support or buffer zone
around a protected area, using such measures as

agroforestry, aquaculture, microllvestock rearing,
increasing wealth
them
among
local people
can give
better tools with which to degrade their
environment, while also potentially attracting

outsiders
who
will
improvements
if
then do the same; and

in
the
livelihoods of
local
wealth generated by
resist
attempts by outsiders
wildlife,
to
they will
destroy
of
let
the
measures
first
lesson
built into
emerges an abundance
of
the family of social investments
known as integrated conservation and development projects [ICDPsl. In this approach, overtly
development-oriented and conservation-oriented
efforts are deployed
280
in
a symmetrical
way
in
the
staff, leafleting
them know what
activities
equipping
surrounding villages
to
the rules are, and reaching out
people to explain ecology and the national
and global importance
of the biodiversity sheltering
the nearby protected area.

If
From
The conservation
protected area, training and
the
to local
in
it.
forestry.
focus on demarcating and patrolling the boundary
enforcement
people are based on their obtaining a share

of the
and community
properly done,
in
a coordinated, respectful
way, with adequate resources and a realistic timeframe, this approach can work quite well. Problems
arise,
however,
when ICDPs achieve much more
development' than 'conservation', when the links
between the aims are not adequately explained,
when
the project cycle leads to a discontinuity
in
shampoo, chocolate,
cream, or cos-
audience shares, however, attractive entertaining
metics are inserted within escapist productions
moving pictures have more impact on the popular
about orangutans apparently
impression
ducts
like
ice
an endless
living in
ram-forest paradise. The combined expenditure
on advertising and publicity for palm
exceeds the money available
for
oil
products far
conserving orang-
utans, and also far outweighs any effect that con-
may
servation broadcasts
some
multinationals should devote

to
Perhaps those
have.
of their profits
great ape conservation' Innocently, or with cul-
pable negligence,
we
(sometimes as shareholders],
tional corporations
Indonesian
we may
politics,
are connected with interna-
and global markets.
be powerless
Individually
change the status guo, but
to
perhaps en masse we could
As Attenborough convincingly reminds

television
and
shows us the wonder
their habitats.
and sustain
producers
of
these primates
the wild' The
in
programs
of 'reality' conservation
that
reveal the conservation crisis have a dilemma: not
home
every viewer will rush
bushmeat trade and the

humans. Those
that do
learn about the
to
shown
to
probably continue to
show and
lovely
in
degradation
feed
makers
to
accuse them
of misleading,
pictures and
or
trust,
when
number
rule breaking by villagers
breakdown
suffer from one or
It
is
more
a rare
of
ICDP
that
even
lying
its
One way forward

into
to take film production
is
communities, making relevant films for local
audiences that would help them
own
value their
wildlife,
to explore
and make
and
a tangible im-
pact on attitudes on the ground. Important pro-
made
gress was
of 11
in
early 2005
when
the
first
batch
award-winning programs on the great apes,
donated
to
the Great Apes Film
Initiative
by
BBC
Worldwide and Granada International, were taken

Brazzaville for local
were crucial
showings there and
for Conservation IFFCI
in this initiative,
and
spread such films more widely
their plan
is to
great ape range
in
states.
Richard Brock
is
their use are distributed
in
ownership, then the whole incentive structure
the success of the project attracts
outsiders to the area.
then
help save these on-
potential to
serious environmental reports; given the relative
to a
will
about the true situation. Television should use
Congo
be watching a mainstream TV
when
its
turn off their televisions and turn on the program
Cameroon. Filmmakers
met by harsh treatment, leading
continuing
Gombe,
viewers
to nothing over the next 10 years,
in
in
channel. The newspapers and radio regularly carry
external support,
its
If
chimpanzees, and Jane Goodall's work are reduced
to
in
National Park,
and endangerment.
The
be few
disturbing
to 'celebrate' the
Gombe
while carefully avoiding the issues of
lUKl are often already aware
of the issues; they are likely to
and unlikely
to
through such an im-
portant but unattractive subject as
Ape Hunters on BBC4
apes
of
killing
sit
than
situation
great apes, filming
us,
Could the media directly repay
source out there
its
the
screen winners but potential real-world losers.
the balance.
tip
of
information about bushmeat. Documentaries will
does not
automatically
change,
according to the
fairly
along
with
values
will
and
behaviors. Villagers who, one year, would cooperate
with gangs of elephant poachers would, the next, be
these problems.
From the second lesson emerges
a family of
reporting the poachers to the police and digging
conservation activities, inspired by the
Communal
waterholes tor the elephants
Areas Management Programme for Indigenous

Resources ICAMPFIREI
This
in
Zimbabwe
in
ment ICBNRMj method has been adapted

locations
ever since,
sanctuaries
in
the
including
Philippines,
bioprospecting programs
ape tourism
is
that
if
of local
in
in
to
many
community
communities and scuba ecotourists
fish
reef-guarding
in
Indonesia,
Costa Rica, and great
Uganda and elsewhere. The concept
wildlife
resources are redefined as an asset
people (whether completely, or shared with
other stakeholders), and
if
For this approach
the 1980s.^^
community-based natural resource manage-
revenue streams from
happened
las
in
Zimbabwe!.
distribution
of
to
work
well, an acceptable
ownership and revenue must be
negotiated and implemented
in
a transparent way;
revenue streams and their distribution must also

offset the
marginal value
to
the system. Even small
person can be enough

are
regular and
to
come
an individual
amounts
change
to
of
of cheating
revenue per
attitudes,
if
they
be expected; future
earnings can then be factored into decisions, such
as whether or not to set a snare

location.
in
a particular
Problems arise when there are disputes
281
World Atlas
creating a long-term social contract

all
concerned.
development
the minds of
In
For integrated conservation and
projects, the costs
Imposed on
local
people by having a nearby protected area are offset

by the benefits offered
the form of livelihood
in
the conservation agency
support by
lor
on behalf
ofl
concerned.
In
systems
like
posed by tolerating
CBNRt^. the costs imearnings
wildlife are offset by
from wild species, mediated by some publicly
NGO
accountable institution such as an
must be
fund. This trade-off
or trust
transparent,
clear,
and trustworthy, and would be
consistent,
fatally
undermined by corruption,
brutality,
favoritism
which range state
(all
factors with
are deeply familiar].
villagers
enterprise,
local
both
In
must have
people
or tribal
kinds of
sufficient
authority to
negotiate freely enough to
and advance
their
own
interests.
protect
There must be
willingness on the part of government to allow them

this authority; this frequently requires a
decentralization or other local
degree
of
empowerment.
STRATEGIC CONSERVATION PRIORITIES

Expert workshops, population and habitat
viability
assessments (PHVAsl, national great ape
survival
GRASP
plans, and the
strategy have
to defining a long list of
occur
in
in
the range states
of the
13. In
priorities
in

Involve
to the precise
circumstances
of
the challenges reviewed
Census training
(top)
and
over the proportion of the earnings going to external
community conservation
stakeholders, for example
awareness labovel
used
Uganda.
in
of
of
armed
a wildlife
may
when
can be summarized as follows:
Is
the
so,
it
will,
The main challenge
282
achieving just and
competence,
institutional
technology transfer as required
in
each case.
science (Including the latest global positioning
in
DRC,
local people
In
the hope
one day, be restored.

to
implementing either
durable arrangements,
systems, geographic Information systems,
and remote sensing)

apes
are,
and
in
to establish
where great
to Identify sites for
tected areas that
type of livelihood-based conservation strategy lies

In
professional and
with resource allocation, capacity building, and
once a revenue stream from
consider wildlife worth keeping
from
that already exist, aiming for high standards of
Surveys and gap analyses. Use conservation
resource has been experienced, attitudes
that revenues
Rehabilitate,
demarcate, and manage the protected areas
of the
Democratic Republic
resource change; as
still
tourism
revenues dries up, as occurred as a result

conflict in the
Congo IDRCl. Even

to that
gorilla
finance parks elsewhere.' or
to
source
when
in
general terms, though, the strategic
Protected area management.

Gordon Miller/IRF
some
area and population, and based on
an understanding
Chapter
to
measures described above and
Chapter U, adapted
of the target
contributed
great apes are to survive
These Interventions would
combination
in
if
summarized
the wild, as
that follow.
all
urgent actions that need
new
pro-
may be needed.
Environmental education. Build local support

for
and participation
in
livelihood
systems that
require ecosystem conservation and are be-
nign to great apes, while also encouraging the
use
of
sanctuaries, zoos, tourism operations,
and protected areas as educational resources

for
Suppression
the
of
ape bushmeat
great
down sources (through enforce-
trade. Close
ment, good management
of protected areas,
provision,
alternative-livelihood
demand
sector cooperation),
them
demarcate and patrol boundaries;
to
and private-
and habitats being conserved; and
the resulting knowledge effectively, putting
In
Money
of
and cooperation across national borders).
microcredit,
easements
cate the initiatives that have already occurred
give
Encourage ex-
apes through partnerships,

journalism, and
investigative
certification,
approaches
adopt
to
consumer and shareholder demand: work

together with
customs
traders,
retailers,
departments, and
society groups
civil
in
and
Understand
initiatives.
vations to conserve and find
up other options
ways
formidable challenges to obtaining

in
in
political
consent
the wealthier parts of the world for the scale of
needed
taxation
fund
to
and
all this,
to organizing
and endowing sustainable financing mechanisms
needed
deliver meaningful
to
and
transparently,
The
local moti-
ways
for local
change
perpetuity).
in
last Is at a particular
(locally,
demands
This
in
will
No such
list
Is
complete
without also noting the need to evaluate every
em-
other resourceful idea that might have
erged already or be about
to
emerge, such as
Is
premium,
them through
taxation
things happen
In
wil-
limited.
is
measures
therefore a need for Interim
make
for tropical
problems are urgent and global
biodiversity
There
biodiversity,
interests.
activities.
who
conservation
themselves appreciate. There are
that they
conservation of forests and
Other
purposes. Those
to participate in
so that the world can have nature reserves and
lingness to solve
own
buy logging concessions or
to
for conservation
people to have the freedom to pursue the
their
to
grants or
inspired leadership and great creativity.
Importing countries.
Community
communities,
great ape populations deserve to be rewarded
agreements.
industries
friendly to great
to finance various
for local
enterprises through
their
Transfrontier cooperation. Reward and repli-
tractive
be needed
will also
compensation
capitalize
Private-sector cooperation.
to
place early-warning and feedback mechanisms.
mechanisms (through targeted enforcement
policy level
of
manage
(through public
and around the Congo Basin, and imple-
to
millions
affecting
lations
forms
ment
programs
people; to survey, study, and report on the popu-
education and alternatives], and distribution
In
protected areas. This will enable
in
sustain outreach
range state populations.
management
national conservation agencies and
teams working
new ways. Nations
that
could
adopt' great ape species, for example, by giving
themselves the task
of
doing whatever
is
needed
to
ensure their survival. With sufficiently persuasive

advocates,
Is
It
possible to imagine an animal-loving
new breakthrough In controlling the Ebola

virus, or a new approach to promoting forests
people
as sacred places.
companies could adopt' World Heritage
the British or Japanese adopting the
like
bonobo as
a national mascot. Meanwhile, private

Sites or
Biosphere Reserves that contain great apes, underIn
doing
these things, there must be a strong
all
emphasis throughout on minimum

local self-sufficiency,
and
The global community
will also
nificantly In
cost,
maximum
effective cost recovery.
these measures
it
need
to invest sig-
writing their
management
permanent source
history of
their'
of
area to sustain and enrich their
public image.
they are to succeed.
In
the final analysis, the limiting factor
This investment need not be wholly financial; self-
whether the peoples
funded volunteers, scientists, and journalists can
apes,
help a great deal.
A range
of
Inexpensive Incentives
can be used by overseas governments

conservation
moval
initiatives.
of trade barriers
Sovereign debt
to
boost
relief
or re-
would free up or increase
range state resources.

Nevertheless, large
needed every year
rain
ecosystems;
amounts
of
cash
be
will
pay
forwages, equipment, training, and running costs
of
of the
many
Is
to
the lack of practical
express our care

Is
one that human

private,
of
in
not
planetary
millions clearly do,
number has nowhere
is
world care about great
and the health
forests,
difference. This
for the foreseeable future to
costs and drawing on a
knowledge about the natural
and
this
go but up. The crucial Issue
means
ways
by which
that will
make
we can
a real
an organizational challenge, and

institutions
(whether public,
governmental, or nongovernmental] are
uniquely equipped to meet.
283
World Atlas
lateral
environmental agreements. These measures
are insufficient, however This
many
partly
is
because so
protected areas are 'paper parks', without the
resources
defend themselves against mounting
to
and partly because so many great ape
threats,
populations and their habitats occur entirely outside

protected areas and are vulnerable to hunting. Wider
and deeper-acting
address the
that
initiatives
underlying causes of habitat destruction are also
seldom forthcoming. The great
required, but are
apes
are, therefore,
Many
decline.
still in
now
conservationists
believe that great
ape conservation requires social mobilization and

a
willingness
public
development that
invest
to
improve their circumstances
ways
in
compatible with great ape survival
especially
that
the wild. Yet
in
These include
Africa.
in
widespread severe poverty and economic and
poster for raising
conservation awareness
In
Cablnda, Angola.
CONCLUSIONS
tical
Conservation activities do not operate

but
variety of external situations, events,
These include war and
vacuum,
and demands.
civil conflict, political
corporate
unrestrained
in
and frequently limited by a
influenced
are
exploitation
of
liuman population increase, the effects
of
and the crippling poverty
in
states,
or to
debt,
to
accept invest-
ments by multinational corporations
that wish to
extract these resources.
Widespread habitat destruction has often been
formerly remote areas. International charities and
interest
community
and concern
in
Efforts to conserve
many
different
locations.
the
Biological
and
Diversity
donors Interested
are able to deploy resources that are paltry and

influence
that
to
multilateral
Heritage
biodiversity
combination
ape conservation, and have undoubtedly helped

preserve certain areas of great ape habitat.
programs and by
local
bilateral official aid
and international NGOs, have
many more
protected areas than
are
designated as globally significant under the multi-
Is
in
Is
with
alliance
nevertheless possible that a
of conditions
consumer groups and

for
being attached
major
to
investigative journalists,
companies
UN
and
that cooperate with certi-
schemes and agree
those of the
to
adopt values such as
Global Compact,
may result
In
more
sustainable use of natural resources.
Meanwhile,
governments, aided and
sector
private
public investments, pressure on corporations from
fication
an enabling environment for great
It
compared with what
trivial
Is
the
governments.
and transfrontier conservation agreements have
encouraged by multilateral and
biodiversity lor great ape] conservation
available
by
rewards
N/leanwhile, national
sustainable develop-
many
strategies and action plans. World Heritage Sites,
to create
in
ment and
UN
World
national
official
them
Intergovern-
have often been led
Convention. The resulting
28/^
service
mid-1970s, the great apes have
activities
aside
to
the inevitable result, complicated for the great apes
environmental agreements, such as the Convention
set
meet
can be aggravated by pressure
many range
agencies or by the UN-sponsored
helped
timber and mineral
natural
have taken many forms and have involved
on
to liquidate
order
in
of
attracted increasing
mental
severe temptation
by hunting pressure facilitated by easier access to
Since the
in
do not encourage long-term planning
and investment. Meanwhile, governments face
current obligations for political expenditure. This
local people.
actors
poli-
armed
nature,
as well as the attitudes, behaviors, and beliefs
international
several cases,
resources
and plans, economic recession,
policies
in
unrest,
disasters and climatic anomalies, other govern-
ment sector
conditions (including,
conflict) that
to
are
many problems and few
the range states have
resources,
sustainable
in
enable local peoples
will
much can
partnership-based
field
be achieved through
projects,
building on the
lessons learned from the educational and other
mechanisms described
above. These
all
involve
complex processes Interacting with environments

that are
themselves complex and dynamic.
In
these
circumstances, the best approach
number
of
Is
adapt to the environment where
is
it
harsh enforcement action, except
avoid
where consensus has been

around the need
process
tently regardless of the details, allowing the

to
adopt a
to
key principles that can be applied consis-
being
nurtured. These key principles are outlined below,
find
in
ways
for this;
built
locally
and
for local residents to
cooperate
unwelcome immigration
excluding
to or
extraction of resources from project areas.
For an educational project:
For projects using community-based natural
3.
of dialog at all
times;
recognize that local people are the actors
and that outside assistance

use
resource management:
maintain respectful, trust-building forms
in
own
their
seek ways
interests;
to turn talk
is
them
for
to
and
build
consensus around an ownership
model
for wildlife resources that includes
local people
and ideas
and has clear consequences
for the division
into action
of
benefits flowing from
as swiftly as possible, to encourage con-
their
fidence and to build 'action competence'.
sufficient benefits to offset the opportunity
use,
so that communities receive
costs incurred;
For programs such as integrated conser-
2.
vation and development projects:
seek
in
and transparency
the distribution
and
itself;
the formula used
ensure that benefits are distributed
development-oriented and conservation-
forms that are acceptable
oriented efforts
communities.
in
the project area;
in
In
to participating
explain clearly and consistently the social

involved
contract'
biodiversity,
in
efforts
conserve
to
and promote improved but
sustainable livelihoods;
achieve symmetry between overtly
to
maintain consistency
to distribute benefits
be alert
to the
sustain-
mechanisms and other
promote
to
of project
them
interventions and build into
means
great apes
this
in
every country where they occur
chapter have been derived, and also
ficult
of
the
it
will
be
to
apply
them
how
in
dif-
effectively unless they
are accompanied by humility, respect, persistence,
and a willingness
continuity;
in
They reveal how the principles explained
wild.
temporary nature
able financing
Chapters 16 and 17 review the circumstances
to learn
and adapt.
FURTHER READING
Adams, W.M.,
Infield,
park. World
Cowlishaw,
Lutz,
E.,
G.,
M. 120031
Development
Who
3^
Is
(II:
on the
gorilla's payroll?
Claims on
tourist
revenue from a Ugandan national
177-190.
Dunbar, R. (20001 Primate Conservation Biology. Chicago University Press, Chicago.
Caldecott, J.O. (19961 Decentralization
Martin, R.B. (19861

of National
and
Biodiversity Conservation. World Bank, Washington, DC.
Communal Areas Management Programme
for Indigenous
Resources ICAMPFIREI. Department
Parks and Wildlife Management, Harare.
AUTHORS
Julian Caldecott,
Box
15.1
UNEP World
Consen/ation Monitoring Centre
Richard Brock, Living Planet Productions
285
World Atlas
Where are
the great apes
and
whose
job
is
it
Ian
great apes
African
mented
belt
of
now
inhabit the
woodland stretching from Senegal

west
are
to
Tanzania
now
in
in
and
the
the east. The orangutans of Asia
restricted to parts of
Map 16a Great ape
Redmond
Despite the detailed
frag-
tropical moist forest
save them?
to
Sumatra and Borneo.
many
there are
apes are or were,

even
differ
as
maps
in
uncertainties about
volume,
where great
Opinions
until recently, found.
how many
to
this
countries
have
still
surviving populations. Their exact distribution
is
distribution in Africa
...S^k.,
WESTERN
SAHARA
ALGERIA
MAURITANIA
Banjul
GAMBIA
GUINEA-
NIGER
"
""Senegal
.
-l
-Bamako
CHAD
,,
_ l^'^P^y
Ouagadougou
BURKJNAFASO
z
Conakry', sierra:
< S a
COTE
Freetown LEONE
,J)'IV01RE
< g *>
assau'WSSAll
,0.N
GWNEA
Monrovia
UBERI4>--
Jâmena
NIGERIA
Ab'uja
CENTRAL AFRICAN
REPUBLIC
Lorn^
Bangui-
RIO ML N
,U&
EQUATORIAI
GUINEA
Kampala
'^
RWANDA
^^vj,
Bi/jufntiiira
bu1m(Ndi
Grea ape distribution
1
<^:,-
'
'<'"^^^DEMOCRATIC
REPUBLIC OF
Bonobo
Chimpanzee
unite6
i
REPUBLIC
pTTANZANLt,
THE CONGO
i'
Gorilla
'
"
if^M
V_ -.ID'S
NGOLA
IN
ZAMBIA^/
Lusaka^
WE
286
...
j'
World Atlas
poorly
documented
in
most cases. Even
is
seldom possible
say when, or even
has been
last individual
apes may
to
live for
killed,
survives and
is
for
years. Moreover,
but
lost
local extinction
is
If
the reasons
can be removed, deliberate
also possible - especially
where
captive individuals, such as those confiscated from

illegal
animal traders, are
site that
need
in
meets the guidelines
Conservation Union. Thus, the
range states
is
of
release
of a
lUCN-The World
number of
great ape
not constant, and can go up as well
as down.
great apes
and woodlands
but are
now
species are found
bonobos
to 21.
The two
gorilla
10 countries and
total of
only one.
in
parts of two
orangutans
restricted
countries, such as
are
in
Southeast Asia, Bornean orangutans are
In
found
in
26 African countries,
of
probably limited
found
and Sumatran
countries,
Indonesia.
to
Rwanda and Uganda,
mainly
some
and are
parks
national
in
In
great apes
the basis of a profitable, carefully regulated tourist

industry,
whereas
in
such as Indonesia
others,
and Cameroon, more apes
outside protected
live
areas than within them, and numbers are spiralling
downwards.
to
In
several countries
100-200 individuals, and
for
numbers are down
some
it
is
too late.
Confirmation of disputed historical reports
Twenty-three countries are known
five
forests
habitat
its
a national border, the possibility
natural re-colonization exists.
reintroduction
the
if,
contiguous with occupied great
ape habitat across

of
it
because individual
more than 50
where a species has been
Chimpanzees may once have inhabited the
those
for
countries where they are reported to be extinct,
now
(Table 16,
Map
more they are reported
within living
memory
Map 16b Great ape
16a,
to
Map
to
16bl,
have
and
in
have been present
difficult,
and even settling questions
distribution faces the perennial
funds for surveys.
Some
of
problem
of
is
current
of limited
the intriguing questions
awaiting a definitive answer include:
(Table 161.
distribution In Southeast Asia

I
Orangutan
T^'.
200
100E
400
600 km
*^
110E
115E
-it^'^'nj.
^'
287
World Atlas
chimpanzees ever
Did
hunters say they used

but
some authors
live
to
dispute
in
Gambia? Old
see chimpanzees^'
this.'
report
question
northwest
map
seasonal movements from across
some
recent
in
the
of
mark over
Togo. This village
in
Burl<ina Faso? There are persistent reports
the Cote d'lvoire border'' and
near
raises
Have resident chimpanzees been extirpated
of at least
chimpanzees
Bunkpurugu on the Ghana-Togo border
is
of the existing
dent chimpanzees survive.' Bance Soumayila,'
Are western lowland gorillas
from Ouagadougou,
convinced that chimpanzees are

in
still
Bas-Fleuve region
is
present
of the
Burkina Faso on the border with Cote
d'lvoire,
near
Comoe
Congo? The
of the
north-
so Togo
profile,
has two possible populations.
biodiversity expert
km
queried record on the
Ghana country
say resi-
from
their extinction
nearly 200
still
found
in
now
the
Democratic Republic
forests are contiguous with
those of Angola's Cabinda province and Congo,
where both western lowland
National Park.
gorillas
and
Table 16 Countries hosting great apes
Genus:
Chimpanzee \Pan]
Species:
Bonobo
Chimpanzee
Subspecies:
western
Nigeria-Cameroon
central
eastern
West Africa
Benin
EXTINCT
Burkina Faso
EXTINCT?
Cote d'lvoire
Gambia
EXTINCT
Ghana
Guinea
Guinea-Bissau
Liberia
Mall
Senegal
Sierra Leone
Togo
EXTINCT?
Central Africa
Angola
Cameroon
Congo
Dem. Republic
X
X
of the
Congo
X?
Equatorial Guinea
Gabon
Nigeria
X
?
East Africa
Burundi
Rwanda
Sudan
Uganda
Zambia
EXTINCT
Southeast Asia
Indonesia
Malaysia
288
World Atlas
chimpanzees are
central
Mayombe
Did
found
still
in
the
Zambia? One
reliable
heard chimpanzees
southernmost shores
is
Zambian
is
the forests
in
of
on the
Lake Tanganyika,
in
in
the 1960s,' but
not usually listed
in
books on
wildlife.
in
rent patchy distribution of
Bornean orangutans
partly a reflection of
human
A glance through
past AO years
to
living
all
across
the past Isee chapter 101?
hunting prac-
the popular literature from the
shows
that people have
been trying
save great apes for decades. Bookshelves are

with the inspiring stories of heroic prima-
tologists, struggling to
save their study animals and
habitats. Conservation organizations have
appeals
to
launched
fund projects. Park guards put their lives
Orangutan [Pongo]
Gorilla IGorilla)
western
Cross River
in
WHO WILL SAVE THE GREAT APES?
filled
Brunei? The cur-
Did orangutans once live
is
north Borneo
in
observer reports having
now Zambia, back
the species
are there ecological reasons which
would have prevented them
chimpanzees ever occur naturally
what
tices, but
Mountains.
Bornean
eastern
western lowland eastern lowland
mountain
northwest
Sumatran
northeast
central
X
X
X
X
X
EXTINCT ?
X
X
289
World Atlas
apes. Educators have
on the
line to protect great
worked
to inspire appreciation of their
many values.
Nevertheless, year after year, the area of viable
to
number
shrinks and the
habitat
This
dvi/indles.
is
great
of
apes
not the result of a concerted effort
eliminate them.
more
is
It
a case of collective
Great apes are being driven
cause people
to extinction
be-
the developed world, the 'haves',
to state-run protected areas,
through community
initiatives,
of
Each
individuals.
the
Important and necessary.
Is
consent
the relevant authorities.
of
where they
among
raw materials
the
and rubber
can only
that
come from
the
support
from plantations that have replaced them. The hard
laws,
economic pressures
And the
habitats.
lift
available
conservation are no match for the massive
for
of
sums
that the relatively small
is
blame
at
or destroy ape
exploit
to
'haves' can hardly point the finger
the 'have nots' - people attempting to
themselves out
poverty by responding to those
of
same economic
pressures
opportunities,
lor
depending on your perspective] - whilst ignoring

their
own
If
role in the equation.
decline
great apes
in
economic community,
is
it
shared by the global
would seem unjust
the job of halting that decline to
governments and people
of the
fall
feel the
government
improve their standard
must be complemented by education, and
efforts
development
by sustainable
habitat. All these activities require
funds and trained
lack both.
Long-term ccnservation success
political
support
will,
view that the survival
of great
the
of
sustainable finance.
sufficient,
apes
live.
budgeted actions
The United
To
initiate
was launched by the

United Nations Environment Programme lUNEPl In
2001 and, joined by UNESCO, was registered as a
Type
Partnership In 2002 at the World Summit
II
kind
governments,
zations
UN
(NGOs),
Johannesburg. This
partnership,
of
of
Identified In
bodies,
civil
whom
bringing together
nongovernmental organiand private sector
society,
concern
of
to
each region.
GRASP, UNEP
range states
It
common
to
Invited the
in
In
govern-
designate a focal point
was requested
be developed
an interest
13.51.
Survival Project (GRASPI
new
Is
and
takes the
wealthier nations to help fund the prioritized,
To help countries work together, the Great
people,
GRASP
of action.
Is
solely to the
have natural populations of great apes
in
Is like
three-legged stool, with three equally Indispensable
the whole of humanity, and has called upon the
- and this includes 16 of the 23 countries
on Sustainable Development
are
personnel, but most of the countries with great apes
annum
Apes
that
initiatives
compatible with the survival of great apes and their
government and begin developing
see Table
to
Thus enforcement
those with a per capita income of under US$800 per
(also
them
helping
is
of living.
ments
still
governments
they understand the purpose behind the
Nations defines 'least developed countries' as
that
those countries
for
range states - those
countries where the great apes
if
and
legs:
the responsibility for the recent dramatic
of
But, in the long run,
live.
make and enforce laws with the support of

people who elect them. People will only give that
oil
ape habitats, from the ground beneath them or
reality
Legally,
therefore, the final responsibility for saving the great
apes rests with the governments
are
projects,
major nongovernmental actions require
from ape habitats carefully enough. Timber and

gold and tantalum, palm
NGO
private sector activities, and, of course, the actions
are not taking the trouble to source raw materials
rattan,
levels -
from International laws
Usually,
negligence.
In
a national plan
that these plans should
consultation with
those with
all
great apes and their habitat, and be
adopted as government
policy.
This task
Is
compli-
cated by the fact that responsibility for great apes
and their habitat
Is
often divided
government departments -
between several
forestry, environment,
tourism, national parks, rural development, etc.
Each country organizes these responsibilities
the
dif-
new administration takes over,

arrangements may change. In addition to this
ferently and,
shifting
mix
when
of relevant
government departments,
case, to ensure the survival of great apes across
GRASP has sought to Involve local and International

NGOs and academics, local communities living In or
GRASP
adjacent to ape habitat, and private sector interests,
Interests, all of
partnership
activities,
is
how
existing
share a
The challenge
their natural range.

to
goal: in this
for the
weave the many conservation
and newly
coherent global strategy
identified.
Into
find the
funds
to
implement
it.
mainly
to
in
tourism and the extractive industries.
Helping 23 nations to develop a coherent
to halt the decline In great
ape numbers, and then - most Importantly - how
290
NATIONAL GREAT APE SURVIVAL PLANS

many
Conservation action can be taken at
cross-sectoral policy to ensure that great apes survive
has proved quite a challenge, and yielded
range
of
results.
Some
countries quickly set up
World Atlas
climate change and
formal national committees, while others took a
the biggest challenge of
less structured approach; several sought funds for
the ensuing disruption to weather patterns and
and one or two had
national workshops,
vegetation zones.
yet to
respond by May 2005. Where the response was
most energetic, the structure
covering
five
The
of the national great
(NGASPl soon took shape,
plan
ape survival
answer
any
such cases, the
detail. In
would be
areas:
Where do
what
is
great apes occur
in
the country and
the current pattern of land use
around their habitat? This immediately

fies the
lii)
mam
What are
area,
region
them
action to counter
What
may
projects,
If
in
virtually every
in
location
still
(v)
is
(iiil
finance
whom,
done
to
It
hoped that the GRASP
is
be greater than the
sum
of
the
from being part
of
the whole rather than
in isolation.
it
of
is
the area of habitat
animals
living
in
left
and the
that habitat, fulfilling
We
can
but hope that in future editions of this atlas, the
country profiles will
will
show
that the 20th
century
decline has been halted, and viable populations of
measurable indicators
of
success
enough
to
new
prices
this end.
the ecological role they evolved to perform.
be
the great apes have survived for the benefit
all
rising
will therefore
number
it?
These plans must be concrete enough
flexible
has provided a global
working towards
each partner brings different
to the task.
for conservation
it?
who
is
GRASP
well
may be decades before we can judge whether

GRASP has succeeded or failed. The bottom line
to halt
best placed to do
the final question
utilized forests, is - to
As with any long-term endeavor, however,
and they are
each area, and what actions need
taken by
developed and implemented. As
for all those
strategy,
working
declining
Having identified the causes of the decline

in
on
knowledge can
individual parts and, moreover, that each partner
been collected - what more do the best

and who
up-to-date
good,
of
gaps
in
whole
where data have
available experts think should be

that decline
surveys
information
the
in
say the
will gain
great ape populations are
despite the activities listed
recommendation
to carry out
managed, sustainably
strengths
truction or overexploitation.
liv)
first
not
stable ape populations at natural densities,
global
law enforcement,
apes and their habitat from des-
that protects
in
is
By working as a partnership within a broadly agreed
local traditions - everything
and
of
framework
the status of current conservation
is
basis
least - patchy. But
effective
if
be taken.
is to
efforts, including legislation,
NGO
each
in
region within
to
must be understood
country, and
filling
effective plans be
identi-
involved? The threats
is
is
known
(i)
the profiles illustrate, progress towards the goal of
the threats to great apes
well vary from
liii]
the
stakeholders.
and who
funds
to find the
states, however,
to question
identified in the following country profiles. Only
and
in
many range
reality in
that even the
remote regions,
lil
all,
to
of
adapt
have
to
of
everyone.
lor failure! but
challenges, such as
commodities from ape habitats
causing 'gold-rush' scenarios
emerging diseases
(e.g. coltan,
le.g. Ebola),
palm
inbreeding
in
oill,
newly
/an
fragmented or reduced populations, and, perhaps
Redmond
GRASP
Chief Consultant.
FURTHER READING
Bailey, N.D., Eves, H.E., Stefan, A., Stein, J.T.,
Bushmeat
Crisis
T.S.,
Apes and Humans:

CITES BWG/IUCN,
eds (20011
BCTF
Collaborative Action Planning Meeting Proceedings.
Task Force, Silver Spring, Maryland. http://www.bushmeat.org/may2G01.htm.

Ttie Ettiics of
Ly, U., Bello, Y.
T.L..
Norton,
B.,
Rowan.
A.,
Stevens,
E.F.,
Arluke,
A.,
eds (2001) Great
Coexistence. Smithsonian Institution Press, Washington, DC.
(20031 Study on Wildlife Legislation
and
Policies in Central African Countries.
http://vTOfw.cite5.org/common/prog/bushmeat/rep_legislation.pdf.
FAO (20031 State of
ttie
World's Forests, http://www.fao.org/forestry/site/scfo/en. Accessed
May 26
2004.
291
World Atlas
Fishpool, L.D.C., Evans, M.I., eds 12001
Important Bird Areas
Publications and BirdLite International,
Forests Monitor 120011 Sold
Down
in
Africa
and Associated Islands.
Priority Sites for Conservation. Pisces
Cambridge and Newbury, UK.
the River The
Need
to
Control Transnational Forestry Operations: a European Case Study.
httpy/www.forestsmonitororg/reports/solddownriver/title.titm. Accessed June 16 2004.
lUCN
119921 Protected Areas of the World:
A Review
of National Systems,
vol.
3: Afrotropical.
lUCN, Gland, Switzerland and
Cambridge, UK.
Kormos,
Boesch, C, Bakarr,
R.,
Plan.
lUCN/SSC Primate
Lee, P.O., Thornback,
J.,
M.I.,
Butynski, T.M., eds 120031 West African Chimpanzees: Status Survey
and Conservation Action
Specialist Group. lUCN. Gland, Switzerland.
Bennett, E.L. 11988) Threatened Primates of Africa. The
tUCN Red Data Book. lUCN, Gland, Switzerland and
Cambridge, UK.
Gates,
J.F.
11996) African Primates: Status Survey
and Conservation Action Plan (Revised
Edition!.
lUCN/SSC Primate
Specialist
Group. lUCN, Gland, Switzerland.
UNDP
120041
Human Development
Report 2004. http://hdr.undp.org/reports/global/2004/. Accessed November 28 2004.
World Bank Group 12005) Countries and Regions. http://wAvw.worldbank.org. Accessed June 17 2005.
World Commission on Protected Areas 12004) World Database on Protected Areas. UNEP-WCMC. http://sea.unep-wcmc.org/wdbpa/
index.htm. Accessed June 17 2005.
292
AFRICA: Angola
Chapter 16
Africa
REPUBLIC OF ANGOLA
Gemma Smith
BACKGROUND AND ECONOMY

The Republic
Angola
of
Zambia
the south,
is
the
to
Terrible
bordered by Namibia
Democratic
the
east,
to
and the Atlantic Ocean
Extending over a total land area of

with a coastline of approximately
is
of
divided
the west.
to
lUb 700 km'
670 km, Angola
One
18 administrative provinces.
into
them, Cabinda province or the Cabinda enclave,
lies to
Congo River and
the north of the
from the rest

territory.
It
is
Angola, and
of
is
separated
km
Angola by about 30
DRC
of
therefore the most northerly part of

Is
bounded by DRC, Congo, and the
Atlantic Ocean. This
Is
the only part of Angola
in
which great apes occur
The history
of
Angola
Intimately
is
bound up
with a long Portuguese colonial presence. This
extended far beyond the period
was
which other
In
European countries dissolved their empires
the
In
1940s, 1950s, and 1960s, because Portugal
was
being run by an ultraconservative military clique

until
'Carnation
the
Introduced
democracy Before
the Portuguese
a
its
which
1974,
of
then, the priority of
government had been
military hold over
included
Revolution'
to
maintain
Timor, rather than to negotiate Independence and

national development of
its
former colonies. The
abrupt collapse of this system

pelled all four territories Into
Portugal pro-
in
independence,
at
which point armed nationalist groups, which had

previously been encouraged by the
China and were
sought
to seize
atmosphere
of a
power.
leftist
In
political
In
USSR and/or
complexion,
War
USA to
the prevailing Cold
of the time, this
provoked the
encourage military Intervention by

Africa in
Angola
Mozambique and Angola,
Guinea-Bissau, and Indonesia
Its allies.
South
military factions
In
East Timor
Is
In all
four
eventually
thus recovering from more than
three decades of warfare,
and then
a civil
first
with the Portuguese,
war between the National Union
for
the Total Independence of Angola lUniao Nacional
para a Independencia Total de Angola, UNITA) and

the Angolan government. This conflict
due
to the
Involvement of Cold
War
USSR-backed Cuban assistance
was
lengthy
interests, with
the Angolan
to
government and US-backed South African assistance
to
UNITA. Major
Involved,
such as
scandal
of 2000, in
in
oil
companies were also
the notorious 'Angolagate'
which the international press
reported that arms-for-oll deals and corruption had

thrived
among
corporate and political institutions
both France and Angola.'^ The end of the Cold
and the introduction
democracy
inclusive
of
in
War
in
South Africa altered these conditions, and the war
ended
In
April 2002, following the death of the long-
term UNITA leader Jonas Savlmbl.'
ended
In all
province.
areas
of the
in
'
Fighting has
country except
in
Cabinda
^^
There are growing signs
possessions, which also
Mozambique, Guinea-Bissau, and East
war and genocide ensued

was
attained.
Republic of the Congo |DRC| to the north and

northeast,
civil
countries, before stable governance
of
economic recovery
Angola, but major problems remain with virtu-
ally
every element of infrastructure and govern-
ment
service
throughout the country, including
communications, roads, and basic education and

health services. Angola exported
billion In
oil
worth US$3-5
2003, about 87 percent of state revenue,
but around 82 percent of the population continue to

live
in
extreme poverty, 42 percent
of
Angolan
children aged five or less are underweight, one child
now
dies of preventable diseases and malnutrition
every three minutes
expectancy
state
is
(I.e.
480 every
44 years. Even
to
day],
and
life
the extent that
revenues are known, somewhere between
293
World Atlas
Map
6.1
in
Angola
12'E
this
country profile
13-E
V.
\
Species
Central
chimpanzee observed before 1983
Western lowland
gorilla
4T
Estimated range
V.Vj' Central chimpanzee
Cf-V.^
Western lowland
gorilla
'v
CONGO
f
i^^K
Buco-Zau
.;'
i
^CABINDA
ANGOLA
1
'\
^^ Massabi
V^[r^K\^DEMOCRATIC
n^^jtS!S&\
REPUBLI
c"'"->..
OF THE CONGO
'^
Cabinda
iW^^SBtSfll^k*^
yMStag^^HjEWKyt
Cabinda
ANGOLA
V*^
1
-.'*^^^gm
REPUBLIC
OF THE CONGO
ANGOLA
_,l..4...l---*^
il
294
"-^
OCEAN
6S
"^
ATLANTIC
12E
'""4>..
CONGO
GABON
ATLANTIC OCEAN
~\
-,
Tir Win ! H^^^^^^^Bnl'

'
AFRICA: Angola
US$1.3 and US$1.7
are missing from the
billion
state coffers.""
difficult
wfiich killed at least
ttie conflict,
to
fialf
people and displaced around another A
a million
number
million. Since the ceasefire in 2002, a large
of internally displaced
persons have returned
to
their places of origin, particularly in rural areas, as
many refugees who had
have
settled
in
neighboring
300
mm (spread over at least
the lower
Congo River and
its
as well as lakes
to
50
In
and are dominated
height,
by tree genera such as Gilletiodendron, Librevillea.
Julbernardia, and Tetraberlinia
Caesalpinloideae). The understory
percent
1998.^
in
crude
the second largest
is
Africa, with
production located offshore
oil
waters
sub-Saharan
in
in
most
oil
of its
the shallow
Cabinda province. Cabinda produces more
of
than half of Angola's

of its foreign
oil
and accounts
for nearly all
exchange earnings.'
With stakes so high,

there are
political
still
tensions
IFLECI, a separatist group,

of
in
is
demanding
a greater
The Angolan government has ruled out
province.
complete independence
2002 announced
its
the
for
province,
autonomy.
of
but
some
operations
Ivtilitary
Cabinda since then, however, suggest that peace

still
killed
living in
refugee
plants.^' The highest annual rainfall
Maiombe
the
in
forest,
and
km'
into
of the
mountainous northeastern
DRC and Congo.

of
tropical
distribution in
This area
moist forest and great ape
camps
in
THREATS
wildlife
of
resources
but
DRC
Development Programme (UNDP)
crops
a
live
is
infant
Massamba,
Cabinda, at the south-
IMayombel
lieved to
tially
at
popu-
figures exist, although residents have re-
forest,'^
be widespread." " On the basis
Cabinda, but there
been thought
Cabinda
poten-
to
may
be more." Gorillas have
be rare there since the 1970s.'
lies within
Mean annual
to protect
chimpanzee was seen
sale
for
In
is
killed."
not traditional
in
The use
of
apes
Cabinda, but
is
occurring, mainly under the influence of ready

in
the neighboring
Meanwhile, the Maiombe forest continues
to
be subjected to a high rate of degradation, mainly
due
to
heavy logging and poaching, for subsistence
and commercial purposes." These observations

suggest that the Angolan populations
ape species may be declining, and
of both great
their distribution
contracting.
it
the part of Angola that

species.
of
was estimated in 1988 that

200-500 chimpanzees could occur in
suitable habitat,
least
Maiombe
where chimpanzees are be-
and
November 2003
In
with the hunter reporting that an adult
male and female had been
and the central chimpanzee [Pan troglodytes
ported sightings of both throughout the
conflict,
conclude that
to
trade,
probably widespread.
DRC and Congo."
definitive
no exception
is
but local information has led the United Nations
markets among soldiers and
gorilla IGoriUa gorilla gorilla)
No
has been seriously
have been deterred by
DISTRIBUTION OF GREAT APES
of their range.
it
affected by conflict and Cabinda

to this. Field studies
known
the richest yet least
In Africa,
The western lowland
lation
interior
the southern
Is
western Africa."
Angola had one
now
western edge
recorded
is
which supports Cabinda's
populations of great apes, and extends across

2 000
as bushmeat
in
the south].
has been replaced with low-density coffee
commercial purposes, the pet
and Congo.
troglodytes] both occur
In
is
increasing
now
parts of Cabinda (particularly
hunting for bushmeat for both subsistence and
population of about 300 000 people, with another
20 000 or more
sparse, and
Is
in
numbers of civilians
by both sides.'' " Cabinda has a
distant, with
being
in
willingness to open talks with
separatist groups, with a view to agreeing
measure
Leguminosae-
(all
Cabinda. The
revenue for the population of the
oil
many
margin
not surprising that
is
it
Front for the Liberation of the Enclave of Cabinda
share
in
It
After Nigeria, Angola
producer
was U2
In
and semideciduous, with
tall
countries.' In 2003, the country's total population
rate of 3 percent per year." Adult literacy
forest
Forests
Guinea-Congo blome occur
the Interior These are
was estimated as
13.5 million people, growing at a
swamp
the coastal plain.
in
characteristic of the
canopy up
and there
tributaries,
are substantial areas of permanent
months
the year) inland. Mangrove communities fringe
of
Recent population estimates are

obtain due to
coast and
the
is
Congo Basin and
most
rainfall is
rich
850
in
mm
LEGISUTION AND CONSERVATION ACTION

International
Angola
is
different
along the
agreements
ratified the
World Heritage Convention
In
1991 (but as yet has no World Heritage Sites], the
UN
Convention
to
Combat
Desertification
the Convention on Biological Diversity
in
in
1997,
1998 (but
295
World Atlas
has not yet completed a national report on imple-
forest responsibilities
mentation
lation
the convention, or a
of
strategy and action
diversity
national
not
is
party
International Trade
Fauna and
the
ICITE5I,
the
2000.
in
Convention
Endangered Species
in
Flora
to
UN
and the
plan],
Framevifork Convention on Climate Change
Angola
bio-
on
Wild
of
Convention
on
Migratory Species, the Convention on Wetlands
formu-
relation to policy
in
and guidance. The provincial government
Cabinda has emergency powers
of
trade
to prohibit
or associated derivatives from Cabinda
in wildlife
within Angola or across
various efforts,
it
is
its
borders." Despite these
reported that wildlife protection
laws are scarcely enforced either inside or outside
of
protected areas, and illegal hunting, harvesting,
International Importance (Ramsarl, or the African
and settlements inside protected areas occur
Convention on the Conservation
regularly.
Natural Resources. Angola
Memorandum
Nature and
of
however, party
to the
Understanding concerning Con-
of
Measures
servation
is,
for
Marine Turtles
Atlantic Coast of Africa, the Treaty of the
of
the
Southern
Endangered
and
wildlife
African grey parrots, bushmeat, ivory,

fairly
openly
in
products,
chimpanzees and other primates,
including infant
markets
etc.,
are sold
Luanda and throughout
in
the country."
African Development Community, and the Protocol
on Shared Watercourse Systems
in
the Southern
African Development Community.
Protected areas
Angola has a long-established system

areas. The
in
Angola dates from 1911,
the establishment of the
deposited.' The
approved
a fund
regulations were
hunting
first
and the creation
1929,
in
parks and reserves
national
of
was mentioned
Regulamento. Legislation
soil,
Fundo de Caca.
Vi/ith
which monies from hunting licenses were
into
of protected
Parque Nacional
national park,
de Caca do lona llona NPI, was established by
National legislation
Wildlife legislation
first
in
the
1936
for the conservation of
fauna, and flora continued to develop, and
in
Regulation No. 2421
of
October 2 1937. Decree
No. 40040 of January 20 1955 provided the
comprehensive nature conservation

the country, covering
and use
of
all
first
legislation for
aspects of conservation
game, and providing
for the
estab-
lishment of national parks, reserves, and controlled
hunting
areas.
also
It
established
Conservation Council, which
laid
Nature
out regulations
was consolidated through Decree No.
governing national parks. Further refinement of
^0040. This legislation formed the basis of the
protected area legislation over recent years has
1955,
It
Hunting Regulations [Regulamento de Caca],

published
1957, and
in
amended
frequently since.
Legislation prohibiting the export of
including
monkeys and
the early 1990s.''
parrots,
first
live
animals,
was introduced
in
'
led to the definition of additional protected area
(outstanding sites
strict
Article 2i of the
Angolan Constitution invests
where
nature reserve
reserve
the state with responsibilities for environmental
which now comprise national park
categories,
(for
nature park
reserve
licensed
(for
public access
is
allowed),
(for total protection],
extraction
only],
partial
regional
nature protection), and special
(for protection of particular species).^'
protection. Since 1998, all biodiversity conservation
National parks are defined by Article 13 of
and protected area management has been gov-
the Hunting Regulations as an area subject to the
erned by the Basic Law of Environment [Lei de
direction
Bases do Ambiente.
for protection, conservation,
responsibility
for
Development
No.
biodiversity
5/98|.
This
moved
from the Forestry
Institute IIFD), within the Ministry of
Agriculture and Rural Development, to the Ministry

of
Environment
(as of
2000 the Ministry
of Fisheries
and Environment and, since 2003, the Ministry
Urban
and Environment)." After
Affairs
the division of responsibilities
is
of
this history,
not yet very clear
animal
more
and control
life
_
fl
of public authorities, reserved
and propagation
of wild
and indigenous vegetation, and further-
for the conservation of objectives of esthetic,
geological, prehistoric, archeological, or other scientific interest, for the benefit
and enjoyment
public* Strict nature reserves,
in
contrast,
of the
'|
were
intended to offer total protection to wild fauna
and
flora (Article 141. Hunting, killing, or capture of
J
]
in
The Forestry Development
practice.
remains
however
in
It
Institute
overall charge of Angola's forest sector,

is
represented
in all
18 administrative
provinces of the country. The National Directorate

of Agriculture
296
and Forest (DNAFI also shares some
animals, or the collection of plants other than for

scientific or
management purposes,
the director general,
is
prohibited
authorized by
in partial
reserves
(Article 15).
Within special reserves (Article 16) the
killing
AFRICA: Angola
species
of certain
is
prohibited, in order to support
Diploma Legislative 88/72 defines
Finally, the
nature park as "an area reserved for the
a regional
and conservation
protection
of
nature,
hunting, fishing, and the collection

of,
in
which
or destruction
of.
wild animals and plants, and the execution of
commercial, or agricultural
industrial,
activities is
priority actions to
Matos
the almost continuous
civil
Improved
ensure
Environmental
Conservation
in
UNDP
2002-2004 by
Norwegian Agency
tion
(NORAD).
for
with support from the
aims
this project
concluded that
1199^1'
war
in
the country since
especially
in
planning, monitoring, evaluating,
environment and
even
in
mammals,
populations of large
Many
large protected areas.
protected areas
lack wardens; poaching, settlement incursion, the

of crops,
cultivation
and the
illegal
collection of
timber and firewood have impacted those sites near
human
population centers.
The Maiombe
unprotected
Angola, yet
in
is
forest
was established
in
1930
was excised
half
reserve
(Alto
area, but
1962.
Maiombe as an important
of
in
when another
Maiombe] was degazetted.
in
centered on Oi AG'S 12 30'E.
watershed
forestry
for
BirdLife International' has identified a

of
virtually
is
the Cacongo Forest Reserve,
purposes." This was originally 650 km^
more than
critical for
taw or practice. The only designated
in
conservation area
which
is
It
bird
is
400 km' part

area
Loeme and Chiloango
the
NBA),
located on the
Rivers,
north northeast of Buco-Zau. and has the greatest
number
the
of
is
of inter-
national environmental conventions.
NORAD
had some
initial
work, from 2000.
request of the Ministry
of
the
justified
improving the long-term
of
the
at
Fisheries and
Environment". The project was
terms
life
in
quality
population of Angola, through stra-
tegically
planned sustainable management
of
natural resources. Key outputs of the project
include a
national environmental
finalized
action plan; the development and implementation of

of a
of
key community-based components
national strategy for the
biological diversity;
on
study
hensive
conservation
launching a compre-
the
state
of
Angola's
environment; establishing an environmental
database; and
environmental
enhancing the capacity
NGOs
to
of
undertake effective
community-based conservation
initiatives,
environmental advocacy, and education.
UNDP/NORAD
to
Following
The avifauna
of
Cabindan provincial government has under-
virtually unstudied, yet its designation
taken to resource the further development
in
forest biome.'"
the
as an IBA by the international conservation com-
of this
munity can only help
servation of the
full
its
Angola that are restricted
species
Guinea-Congo
the area
the nation's
of
implementation
of
forest in Cabinda
strengthen
to
and manage natural biological resources,
protected areas and conservation efforts, particuto
in
Development Coopera-
and reporting on the state
relating
and
Diversity
Angola (project ANG/02/005). Implemented
1974-1975 has had significant impact on Angolan
larly
Planning
Biological
of
conservation
its
identified.
national capacity to protect the environment
prohibited or conditioned.'^
Huntley and
and
and sustainable use
their conservation."
to
strengthen the case for
its
project,
the
work. Plans for the study and con-
Maiombe
forest
and
its
ape
protection.
The Friends
United Nations support
UNDP
is
of Nature
club of Ganda-Cango,
supporting the Angolan government
in
seeking ways of sustainably protecting the environ-
ment and managing natural
biological resources."
Angola.
a
Members make
commitment
not to eat
bushmeat.
Key projects are described below.
Development
of a national biodiversity stra-
tegy and action plan. Through this project,
which
is
Facility
funded by the Global Environment

(GEFI
November
2004.
UNDP
and
it
is
and started
hoped that the status
in
of
the biodiversity of Angola will be defined, the
pressures to which
it
is
exposed documented.
297
World Atlas
populations are being
developed.
Initiated
through) legislative
developments
and provincial
the proposal includes the
level,
designation of a
new
at national
protected area and a
unique ecosystems
be
While wartime
lost.
some
Angola's especially rich crop
population,
chimpanzees and
scale introduction of imported commercial seeds
wildlife including
based on the involvement
is
It
communities
local
in
developing alternatives
species, and education and public
campaigns. At the request
UNDP
helped
initiative to establish a
task force for the joint
Maiombe
awareness
the Ministry of
of
Fisheries and Environment,
coordinate an
of
and
forest
its
flora
it
to
regional
protection
the three countries that share
of
the
communities, the
approved by
all
process."
The
in
awareness, and alternative livelihoods are the basic
requirements for the sustainable protection

diversity
in
much more
has been
tradition.
which has no strong ape bushmeat
Hunting for subsistence can be resolved
mainly through the combination
Maiombe area
consultation process
armed
with resident communities and, within the

forces,
the development of voluntary 'friends of
whose mem-
nature' clubs lamigos da natureza],
bers have agreed not to eat bushmeat except
extreme survival situations." The association

operators
CABGOC
established the
Protocol with the Angolan
2002, and provides modest support
in
Maiombe
to the
of oil
Cabinda, headed by ChevronTexaco,
in
government
in
There are several immediate
awareness campaign among
soldiers, police, resi-
dent communities, and others within the forest, to
ensure their cooperation

forest
and
economic status
means
is
of
The devastated sociocommunities, however,
local
that an education
the protection of the
In
biodiversity.
its
and awareness campaign
not likely to be sufficient on
accompanied by measures
to
its
own, but must be
encourage and enable
Such
and sustainable
of the objectives of the
There
ment, with the additional aspects
virtually
no enforcement
of
environmental
Angola, so the overwhelming priority for
mental capacity
for
must be
building funda-
especially
in
main areas
in
throughout Cabinda, and
and around the Maiombe
of
be addressed
efforts
level
forest. Six
environmental concern also need
to
at the national level in parallel with
Cabinda. These are deforestation,
soil
communities, and
of
Agriculture,
employ the
by
biological
of
fish
stocks;
by the petroleum industry; the loss of

diversity;
and
poor
environmental
sanitation. '^ Civil conflict has resulted in the break-
down
to
of the protected
area system established prior
independence (covering
national territory], and there
6.5
percent
is
a risk that
of
the
some
and Environment
Fisheries,
will
biodiversity advisor originally funded
organization,
also working
is
Gremio ABC.
Priorities
include the
recruitment of a law enforcement unit of rangers

(foca/sl,
pollution
visibility
UNDP/NORAD. The department
provide
energy sources;" the depletion
assuming
with a provincial environmental nongovernmental
impoverishment, erosion, and desertification;' the
poor households on traditional
of
From August 2005, Cabinda's Department
dependence
of
Cablndan provincial govern-
social responsibility. Improving relationships with
enforcement and education
governmental, nongovernmental organization,
and community
framework
efforts are within the
FUTURE CONSERVATION STRATEGIES

is
priorities,^'
including the further development of an extensive
livelihoods.
in
developing
of
awareness campaigns.
local people to achieve alternative
forest area.
is
may be
destructive than subsistence hunting
this area,
in
of bio-
Cabinda. Effective law enforcement
(Angola, DRC,
initiative
the
to identify potentially
viable populations. Education, enforcement, public
alternatives and
include an awareness and
great apes include a
of
Cabinda
of
essential to halt commercial hunting, which
three governments.
Other conservation efforts
needs
priority
thorough census
and fauna by
and Congo). Mainly based on the participation

of local
during the postconflict agricultural rehabilitation
unsustainable use of forest
to the currently
at
plant
genetic resources could be threatened by the large-
gorillas.
law
protection for
both plant and animal species, due to rural de-
separate sanctuary for the rehabilitation of
orphaned
298
will
conditions have also provided
of
alternative livelihoods,
ownership. There
first
In
mainly from resident communities,
some
group
of
Is
an urgent need
others.
In
2003,
it
budget allocated under the

for
to train a
rangers that could then take part
establishing the law enforcement unit and
training
the
training
contribute to a
to
and a feeling
of
first
was agreed
In
that the
UNDP/NORAD
project
rangers would be used
to
national course for rangers by
"
AFRICA: Burundi
the Southern African Wildlife College,
NP, including places for
six trainees
conservation
Finally, a transfrontier
is
response
smuggling
timber,
wildlife,
of
involve all three
to
countries that share the forest: Angola, Congo, and
UNDP
DRC.
initiative
development
has produced
of
such an
proposal tor the
initiative,
and
has been
it
distributed to the international scientific and donor
harvesting and cross-border
the
to
would need
derivatives. This
for the protection of the fvlaiombe forest,
needed
in
Kissama
in
from Cabinda.
and associated
community."
FURTHER READING
FAO (2001 Forestry Outlook Studies
1
for Africa
IFOSAI - Angola Country Study FOSA Working Paper - FOSA/WP/02.
FAO, Rome. ttp://ttp.fao.org/docrep/fao/003/X6772E/X6772E00.pdf. Accessed

Huntley, B.J., ed. 11994) Botanical Diversity
in
Southern
lUCN 11992) Angola: Environmental Status Quo Assessment Report

Southern
May
19 2005.
Africa. National Botanical Institute, Pretoria,
- Executive
Summary. lUCN Regional
Office for
Africa, Harare.
lUCN 119921 Peace
in
Angola.
lUCN
Bulletin 23.
Ministry of Fisheries and Environment 12002) Report to the
Convention
Combat
to
eng.pdf. Accessed
May
Committee
for the
Review of the Implementation of the
Desertification. Luanda. http://wAA.unccd.int/cop/report5/africa/national/2002/angola-
19 2005.
MAP DATA SOURCES

Map
Great apes data are based on the following source:
16.1
Butynski, T.M. (20011 Africa's great apes.
Stevens,
E.F.,
Arluke,
A., eds.
In:
T.S.,
T.L.,
Norton,
B.,
Rowan.
A.,

With additional data by personal communication from Ron,
T.
120031.
ACKNOWLEDGMENTS
Many thanks
the
to
Maiombe
Tamar Ron [UNDP]
for her valuable
comments on
the draft of this section, and for information about
forest.
AUTHOR
Gemma
Smith,
UNEP World
REPUBLIC OF BURUNDI
Gemma Smith

Located
is
in
to the north, by the
central Africa, the Republic of Burundi
a small, steep country that rises
from the east-
ern shore of Lake Tanganyika. The land area
25 650
km^
section
of
000
mountain
km^ including Burundi's
or 27 834
the
lake.
along
the
range,
Elevations only
lake's
the
is
fall
below
shores 1773 ml. A
Congo-Nile
Ridge,
that
United Republic of Tanzania to
the south and east, and by the Democratic Republic

of the
Congo IDRCl
lation
was estimated
up
to
areas.
to the
west.'" Burundi's popu-
at 6.2 million in 2004,^ with
300 people per square kilometer in some

" The population is growing at an annual
''^^
rate of 1.9 percent.^'
groups: the
It
includes three main ethnic
Hutu lover 83 percent), the Tutsi
than 15 percent), and the Twa H percent!.
reaches 2 670 m, runs roughly north-south along
(less
the western boundary, providing the highest land
Approximately 52 percent
in
the country. Most of the country comprises
plateaus and rolling
hills.
It
is
flat
bordered by Rwanda
age
of
inhabitants over the
of 15 are literate.'
Burundi's economic situation has varied
in
299
World Atlas
Map
16.2
Chimpanzee
distribution in Burundi
20
10
30
this
country profile
50 km
40
r'
X'
4-
/M
RWANDA
/
Lac
Cohoha
Slid
Ri
1
1
'X-^
Kibira
NP
1
Muyinâ
3"S
a-'si
Karusi
Cankuzo
Ruvubu
_^-'^
NP
UNITED REPUBLIC
DEMOCRAft-IC
REPUBLIC?' OF
THE CONtÔ
4'S
Rumonge FR
Kigwena FR
i"
Vyanda FR
Mukungu.,
Mabanda/ NyanzaLake PL
''.^
Rukamabasi PL
'
Species
Eastern chimpanzee observed 1996-2003
Eastern chimpanzee observed after 1983
Eastern chimpanzee observed before 1983
^^\4
Eastem chimpanzee
Eastern lowland
locally extinct since ca.
gorilla
Estimated range
^.\j> Eastem chimpanzee
Eastern lowland
300
gorilla
1940
AFRICA: Burundi
recent years due to
disturbances and
political
2002, the gross domestic product (GDPI
conflict. In
was US$719
and
nnillion
IGNI! per person
gross national income
ttie
was US$110." The economy
is
based on subsistence agriculture."' " with 90 percent of the population engaged
farming.' Coffee
in
is
Burundi. Relatively
known about Burundi's
little is
chimpanzees, but 300-400 individuals are thought

to
Surveys
exist there.
and 30-50
reserves." " In
IMP]
1987 found two small
in
200-250
populations;
Rumonge and
the
in
based
1992,
Park
National
Kibira
in
on
Bururi
surveys
by
made for popu-
Burundi's main export, while principal food crops
Trenchard," further estimates were
are cassava, bananas, maize, sorghum, and beans.
lations
Livestock are abundant and there
Mabanda/Nyanza-Lake and Mukungu-Rukamabasi
pressure
soil
many
in
erosion
heavy grazing
locations, leading to
damaging
The varied topography,
number
soil,
Most
and climate sup-
of vegetation types."
the natural vegetation
is
mosaic
of
the Vyanda
in
Most
of
East African
altitude of
country.^
600-2 666
tiguous with
of
afromontane
by Brachystegia and Julbernardia (both
nated
montane
of the
Nyungwe
has been degraded by grazing and
in
sum
extended over
bamboo species occur
country.'
In
however, only an estimated
2000,
940 km' remained under
percent of total
forest, 3.7
about 16 percent
from legal and
of
leading to rapid
Burundi gained independence from Belgium

July 1962. Since then
it
has been plagued by
who had
tension between the Tutsi minority,

ditionally ruled the country,
intractable conflicts in Africa.
elections occurred
in
Hutu head
1993,
tra-
and the Hutu majority,
and Burundi has been the scene
their first
one
of
The
first
of the
most
in
of state,
Key tree species
1993,
forest,
gummifera
at
of the
300
layer and trees
altitude, while
Some
900-2 300 m.
It
is
estimated that
park comprises primary
mostly montane." Since the crisis
however,
much damage has occurred
democratic
bush
fires,
illegal
timber harvesting, agricultural
encroachment, goldpanning, poaching, collection
of
medicinal plants, and charcoal production.'

Bururi
Forest Nature Reserve
southwest Burundi,
to
is
located
in
the west of the town of Bururi,
with an altitudinal range of
900-2 307
and an
Melchior Ndadaye,
for
Table 16.1 Estimated chimpanzee populations
in
Burundi, 1992
Burundi (Frodebul. Within months
Ndadaye had been assassinated,
setting the scene
which an
tor years of Hutu-Tutsi violence in
mated 300 000 people have been

conflict, in April
a Hutu,
succeeded Pierre Buyoya, a
10
2003, Domitien Ndayizeye,

Tutsi,
Park, reserve
Vegetation
Area Ikm'l
as head
Burundi's three year transitional powersharing
government.
Number
of
chimpanzees
esti-
killed. After
years of
of
of the
when Burundians chose
and a parliament dominated by the Hutu Front
Democracy
300 km'.
through uncontrolled destruction by armed gangs,
deforestation.'
in
at 2
trees grow as high as 50 m.
evergreen
logging,
each major part
A herbaceous
(Meliaceae)."
land area. The forests are under intense pressure

illegal
con-
and Entandrophragma excei-
(both Leguminosae),
occur on rocky soils even
of the
is
Rwanda, forming
I'tioesti].
natural and planted forests of Burundi together
000 km', about 8 percent
north-south
include Newtonia buchananii, Albizia
farming. Prior to the conflicts of the 1990s, the
an
Endangered species LHoest's
sent, including the
vegetation
at
park.^ Several other primate species are also pre-
guenon iCercopithecus
of the natural
the
forest block of approximately
Leguminosae-Caesalpinioideael exists along the
Much
forest in
Chimpanzees are found
Kibira
the northwest of the
in
Congo-Nile Ridge, and
southeast border, with small patches of transitional

rain forest in the northwest.
stretches along
It
land with abundant Acacia trees ILeguminosae-
Miombo woodland domi-
live in
NP, which covers approximately 403 km'
mountain range
Mimosaceael, with large areas
16.11.
Burundi's chimpanzees
of
evergreen bushland and secondary wooded grass-
vegetation to the west.
Reserve and the
Forest
Protected Landscapes (see Table
this high rainfall area."
in
port a large
is
Kibira
NP
Bururi Forest Nature Reserve
Afromontane
forest
403
200-250
Afromontane
forest
33
30-50
Miombo woodland
10-15
Vyanda Forest Reserve
Miombo woodland
45
50-68
Mabanda/Nyanza-Lake
Gallery forest
85
15-20
Rumonqe
Forest Reserve
and Mukungu-Rukamabasi
Protected Landscapes

The
eastern
schweinfurthii]
chimpanzee
is
[Pan
troglodytes
the only great ape to occur
Total
305-403
in
301
World Atlas
Kibira
NP
much
suffered
land conversion and forest
degradation during the 1990s.' Habitat fragmentparticularly affects the chimpanzees of

Rumonge, Vyanda, and tvlabanda/Nyanza-Lake,
which now move from one small forest block to
ation
another Along the way, they come
humans and
with
sometimes
are
into close contact

killed.'
LEGISLATION AND CONSERVATION ACTION

International
Burundi
The impact
of gold
Diversity in 1992;
mining at Mabayi, near

the
Rwandan
agreements
signed
ratified
jecttroy Citegelse
area of 33 km'.
contains about 16 km' of semi-
It
evergreen forest, and receives an average annual

rainfall of
200-2 400
tree species fiave

of
mm.
Despite
been recorded
its
small
Myrianthus ICecropiaceael
size,
93
species
tiere, witfi
Strombosia
and
(Olacaceael being dominant, and ttiose of Newtonia
Tabernaemontana (Apocynaceael,
(Leguminosael,
and Entandrophragma being common."

Vyanda Forest Reserve
Zambezian
Desertification and the
Iboth
Species
to the
IRamsar]
savanna vegetation
in
fias
an area
dominantly composed
forest.
of
of
some
6 km'.
uniform
is
also
is
It
pre-
Brachystegia
Other primate species present include olive
baboons [Papio anubis]
and
monkeys
vervet
[Chlorocebus aethiops].
The
protected
in
in
the
Endangered
it
Con-
ratified the
has
ratified
tvlan
it
19821,
World
the
Heritage
and participates
currently has no designated
MAB
sites
or properties inscribed on the World Heritage

It
is
not
in
and Biosphere IMABI Programme,
the Convention
party to
List.
on Migratory
Species, despite the presence of several species

listed by the
regional
convention within
biodiversity
treaties
Other
its territory.
and agreements
of
Mabanda/
Nature and
of
Natural Resources.
the south of the country at an altitude ranging
between 900 and

85 km'
in
600 m, and
total
approximately
area. Natural vegetation covers about
made up of open forests dominated

Brachystegia, wooded savannas, grassland
37 km', and
by
ratified
party to
which Burundi has signed include the African
landscapes
Nyanza-Lake and Ivlukungu-Rukamabasi are located
1988, and
in
Convention (May 19
although
Forest Reserve
situated in western Burundi, at an altitude of about
850 m, and
also a
2002.
Burundi
common
Rumonge
Combat
to
Wild Fauna and Flora (CITESI, acceding
of
convention
UNESCO's
DRC.^
is
from East Africa and lowland evergreen forest

in
Burundi
19971.
in
bio-
been
Convention
on Climate Change have been signed and
habitat, as well as a
flora, including drier
have
plan
UN Convention
UN Framework
Both the
'
action
vention on Wetlands of International Importance
situated nearby, and
is
comprises miombo woodland
strategy and
developed.'
Biological
and a national
of the convention,
diversity
on
came into force in 1993 and was

Two national reports on imple-
1997.
in
mentation
border.
Convention
the
it
is
savannas, submontane gallery forests, and low

grassland.
study of the fauna of these protected
The
National pour lEnvironnement et
Institut
Conservation de
is
la
Nature IINECN], created
responsible for the
management
parks, nature reserves, and natural
Part
of
areas remains to be made, but olive baboons and
Territoire,
aardvarks lOrycferopus afer] are found here.^
of
the
Ministere
de
de lEnvironnement
of
in
la
1980,
national
monuments.
I'Amenagement du
et
Tourisme (Ministry
Land Planning, Environment, and Tourism),
it
is
also responsible for organizing and implementing
THREATS
biodiversity
Chimpanzees are vulnerable
to
hunting, logging,
and forest clearance, so they are assumed

decline, but the rate of loss
of
is
unknown. The
be
in
effects
warfare and conflict have further exacerbated
the impacts of habitat loss." The
302
to
montane
forests of
ties,
assessments and conservation
activi-
for organizing public conservation education
activities, for
making proposals
tor
new
protected
areas, and for ensuring that tourist sites are used
sustainably" INECN
and
is
also the CITES
management
scientific authority for Burundi.
AFRICA: Burundi
There
no national law that specifically
is
protects chimpanzees, although export of primates
As chimpanzees are thought
is illegal."
be res-
to
tricted to protected areas, their current populations
assumed
Currently, four types of national protected area
are designated
Burundi: national parks {pares
in
nationaux], natural reserves [reserves naturelles],
monuments [monuments
natural
and
naturels],
be sufficiently protected from
protected
landscapes [paysages proteges]."'
hunting by the protected area legislation. The 1971
Fourteen
national
are widely
to
measures
regulation on hunting and protection
for
certain animal species includes 12 articles relating

to
hunting rights, the special permits required for
hunting,
methods
listing of
protected species. Other relevant national
of
hunting prohibited, and the
conservation legislation
"
outlined below.'
is
protected
ment
monuments
Iseven], natural
landscapes
1
(four).
which
habitation
areas are prohibited, as
protected
exploitation within
Law
and
km
land
is
provides for establishing protection forests,

reserves, and
areas,
reforestation
is
administrated by the Forest Service, and aims

to protect soils
and conserve animal and plant
species under threat within forest reserves
and protection
Law
Burundi's key legislation to allow the
as an
is
man-
protection of the environment
integral
part
of
Environment Strategy.
in
In
Ramsar
Tanganyika.
Conservation projects
1992,
INECN
in
Goodall Institute
collaboration with the Jane
(JGI) initiated the Kibira
anzees Project - but
war
WWF-The
2002,
In
was
this
by the
Conservation
Global
Organization started to support work
aiming
Chimp-
derailed
Kibira NP,
in
encroachment and destruction
to avert
is
carried out through the
ecoregional project for the Albertine
and
undertaken with Burundi's parks
being
is
WWF
Forests,
Rift
authority and two local nongovernmental organi-
zations (NGOs): the Association Burundaise pour
63 articles, the code
Protection des Oiseaux (Burundi's BirdLife
ABO) and the Organisation pour
vironmental impact assessment procedures,
I'Environnement au Burundi (ODEB). The project
protection and development of natural resour-
funded by the MacArthur Foundation.
forms
cultural
human
of pollution,
heritage,
environ-
control
of
all
of Kibira
NP and
resource
four
vation
of
ABO and
and Kigwena), and defines their
nizations and
media
objectives and
the
mode
of
ing
communities
in
economic and
ecological benefits that the forest offers.'
forest users of the
The project supports and reinforces the

of
I'Association
Burundi
is
one
of the
populations affected by
few African countries that did
have an established protected area system
during colonial times, and
it
was
not until
formal protected area legislation

tence here.
came
moting
efforts
d'Encadrement, de Production
de Vulgarisation (AEPV-DUFASHANYEJ
Protected areas
activities to
In
develop an under-
to
the sites. The protected landscapes were also
boundaries had
to
conser-
Burundi have been organiz-
standing
not yet been delineated.^
in
income or benefits."
protection and conservation of the flora within
identified here, but the precise
aims
other environmental orga-
awareness campaigns
among
improve
to
and
is
provides
It
while exploring and promoting
sources
recent times,
management
Defense de
la
park authorities
management,
involve local
activities,
alternative
to
Rumonge,
reserves (Bururi,
natural forest
Vyanda,
NP, Rusizi
support
natural
empower and
and penal provisions.
Decree No. 100/007 (20001 delineates the

boundaries
training
la
affiliate,
addresses administrative responsibilities, en-
ment and
of
the area as the country returned to peace. This
Burundi's National
ces including biodiversity, the
not
designated
Site
2002, and covering 10 km' of the shore of Lake
partnership work
forests.
No. 10 (2000), the Environment Code,
agement and
land
area.'""
In
No. 1/02 (1985), the Forest Code, which
forest
over
of
boundaries.
of the
of the total
To date Burundi has designated one inter-
and nature reserves,
hunting
and protected
(one],
Together they extend
277 km^ and cover A.6 percent
'"'
1/06 (1980) provides for the establish-
of national parl<s
within
"
designated; national parks (two), natural reserves
national protected area, a
Law No.
"
been
have
areas
war
It
does
et
to help local
this by pro-
generate income, supporting
forestry and small plantations around the parks,
980 that
and by raising awareness among the people and
into exis-
the administrative and military authorities about
the Parks for Peace concept,
in
order to transform
303
World Atlas
this
park
into
be respected
that will
site
by
The status
everyone, even during periods of conflict.'
lUCN-The World Conservation Union and
the
in
phase
of the
includes the
NP
Kibira
NP
Virunga
Parks
initiative
Peace
project,
NP
the Volcanoes
aims
which
nerability of the eastern
The establishment
as well as
Burundi,
in
DRC and
in
Rwanda. The
for
to resolve conflicts
and
is
uses
Conservation Society (WCS)

region through
its
Albertine
is
also working
Rift
in
in
ted the
Jane Goodall
1994,
relocate the 20
center
in
chimpanzees from
Bujumbura, Burundi,
orphans had mostly originated
in
It
A few
need
bio-
undertake
to
to
determine pre-
essential to continue raising

its
awareness
current degradation.
national organizations are already doing this,
efforts of international NGOs [such as WWF and

WCSI should continue and their efforts and their
Kenya. These
DRC."'
is a
but further efforts are needed. The constructive
to
rehabilitation
its
to
IS
about the environment and
permission
Institute to request
assessments, there
have been historically recorded,
promp-
from the Burundian and Kenyan governments
be required, as
sence and numbers.
Burundi, primarily due to the relatively

in
bio-
surveys within protected areas where chimpanzees
the
Programme.
unstable political situation, which,
likely to
is
As well as undertaking more general

diversity
There are no sanctuaries or reintroduction

centers
new environmental and
tected areas, poaching, and timber harvesting.
reduce conflicts and the
protected areas. The Wildlife
in
may
has been developed, additional
capacity to implement this
threat of poaching, logging, mining, and other destructive land
to extinction.
protected areas
well as tight controls on encroachment into pro-
as well as providing training and support for

is to
essential. Although
diversity legislation
partnerships and dialogs at national and local
survey work. The aim
chimpanzee
of additional
be required. Improved environmental governance
in
impacts upon protected areas through enhancing
levels,
parks and forest reserves
regarding the increasing isolation and resulting vul-
Burundian government have completed an orientation
of national
Burundi urgently needs assessment, especially
'""
effectiveness should be monitored.
FURTHER READING
Habonimana,
A. [20031
Convention
on
The Magnificent Kibira Parl< Turned into a Land of Devastation. Submitted

www.unccd.int/publicinfo/localcommunities/burundi-eng.pdt.
Desertification,
to the
UN
Accessed
September 27 2004.
Powzyk,
J.
119881 Tracking Wild Ctiimpanzees in Kibira National Park. Lothrop. Lee and Shepard,
New
York.
MAP DATA SOURCES

Map
16.2 Great
apes data are based on the following source:
Butynski,T.M. 120011 Africa's great apes.

Stevens,
E.F.,
Arluke,
A.,
In:
Beck, B.B., Stoinski.TS.. Hutchins, M., Maple, TL., Norton,
eds, Great Apes
and Humans: The
Ettiics of
B.,
Rowan,
A.,
Coexistence. Smithsonian Institution Press,

With additional data from:
INECN
11992) Projet
Chimpanzees de
la Kibira -
Proposal submitted to the Jane Goodall
MINITERE
1200A)
Jane Goodall Chimpanzees. Conservation and Research
Rwanda's National Great Apes Survival Plan 2004-2009.
Forestry, Water,
Project.
Institute.
and Natural Resources, Republic
of
Final draft. Ministry of Lands, Environment,
Rwanda.
ACKNOWLEDGMENTS
Many thanks
to Geoffroy Citegetse lABOl,
reviewer for their valuable
comments on
Adrian Habonimana lABOl, Jean Rushemeza IINECNI, and an anonymous

the draft of this section. Thanks also to Brigid Barry ITropical Biology
Association! for editorial assistance.
AUTHOR
Gemma
30<l
Smith,
UNEP World
AFRICA: Cameroon
REPUBLIC OF CAMEROON
Caldecoh
Patrice Taah Ngalla, Lera Miles, and Julian

The Republic
Africa on
tfie
Chad
west,
Cameroon
of
Cameroon chimpanzee and Cross

located
is
in
Central
Gulf of Guinea, with Nigeria to the
Republic ICARl
to
Central African
the
north,
the
to
the east, and Congo, Gabon, and
Equatorial Guinea to the south.
in
the
in
Western
the highlands of the north-
around Lake Chad
In
range
the extreme northwest Inow
Mount Cameroon
desiccated], and the
of forested
mountains
In
the southwest.
was growing
2003, which
in
about 2 percent per year'^ Approximately half

in
most
of the rural population residing In the
gross
national
of the country.
Trinationale de
south
Income (GNII per person was
US$560, and the gross domestic product (GDPl

the country
was US$9.A
billion," 9
for
percent of which
NPl,
300 km", with buffer zones
Congo
and
NPI,
of
the
called
collectively
Sangha. The core area
la
to live
common
(Lac Lobeke National Park
CAR (Dzangha-Ndoki
at
2002, the
In
Cameroon
frontiers of
(Nouabale-Ndoki
live
Cameroon.^' These
in
000 gorillas thought
of the 10
of
urban areas such as Douala and Yaounde, with
and central regions
some
that there are 15 000
is
a triangle of protected areas on the
(NP)I,
Cameroon had an estimated population

15.7 million people
to
lovyrland gorillas
The most recent estimate
southwest,
the
In
include
somewhat
and central chimpanzees are widespread
the south of the Sanaga River (Map 16.3al.
west and central provinces, a seasonal marshland

In
[Mandrillus leucophaeus]. Western lowland
drill
gorillas
moist
coastal plain
savanna woodlands
the
covers approxi-
It
mately 475 UUQ V.m, with forested lowlands

south,
River gorilla,
as well as other restricted-range primates such as
is
around
21 000
kml"
Other recorded locations include the Campo-Ma'an
NP,
Forest
Management Unit
003, Mengame
dAmenagementl
Dja Wildlife
Forestlere
(Unite
Gorilla Sanctuary,
Reserve (also known as Dja Faunal
Reserve, and designated as a Biosphere Reserve
and a World Heritage
Site],
the
Abong-Mbang
Forest
derived from timber products." The national eco-
Reserve, Deng Deng Forest Reserve, a small pocket
nomy has
of forest
a strong
dependence on
oil,
timber, and
outside
Nanga Eboko, and the Southeast
cocoa exports; the major national Industries are the
Forest Technical Operational Unit [Unite Technique
production of timber and textiles.
Operationnellel, which covers
The present country was formed

through the merger
of
the former French
in
Cameroon
with the English-speaking southwest, which
then administered by the
UK
1961
was
(and had been part of
German colony of Kamerun until the First

World War). Compared to other countries in the
the larger
region,
it
has experienced relative
stability (apart
to the
border with
thought
Edea
to
CAR and Congo." "
Mengame
Sanctuary
Gorilla
at a
Cross River gorillas

In
Cameroon, the Cross River
Overview
Reserve, the Mbulu Hills

gorilla [Goritla gorilla gorilla],
(G. g. diehli],
central chimpan-
[Pan troglodytes troglodytes], and Nigeria-
Cameroon. This
[P.
diversity
t.
is
vellerosus] all occur

a
result of the
In
bio-
geographically unique, transitional nature of the
Cross-Sanaga area
of
southwest Cameroon and
southeast Nigeria. This region contains the Nigeria-
200 western
higher density further from
the
Cameroon chimpanzee
the
and settlements."'^'
villages
zee
in
greater
and the
and timber Industries.
Cross River gorilla
lowland gorillas and 200 central chimpanzees. The
human
The western lowland
the Douala-
found
population of apes than expected -
animals were found
of Infrastructure, agriculture,
Gorillas are
In
A 2002-2003 survey
Wildlife Reserve."
Peninsula, ongoing since 19921, allowing the steady
oil
eastern
of the
be rare and possibly extinct
from a border dispute with Nigeria over the Bakassi
development
most
province of Cameroon, from the Lomle-Batouri axis
Takamanda
other highland areas

of
gorilla is confined to
Forest Reserve, the
Mone
Community
of forest to the
Forest
Forest,
and
east and south
these,'"" as well as the nearby forests
in
Nigeria.
These are the northernmost and westernmost
gorilla
populations, separated from the range of the western
lowland gorilla by
Cameroon and
of
largely
some 200
Nigeria
is
km.''
The range over
limited to at
most 200
km''
unconnected forest fragments, spread
over a total area of
at least 2
500 kml''
'^
There are
305
World Atlas
Map
16.3a
Chimpanzee
Cameroon
distribution in
country profile
this
Confirmed range
Species
996-2003
Central chimpanzee observed
Central chimpanzee observed after 1983
Central chimpanzee observed before 1983
Central chimpanzee presence alleged
^.\j- Central chimpanzee
Nigeria-Cameroon chimpanzee observed 1996 -2003
i-lj- Nigeria-Cameroon chimpanzee
Nigeria-Cameroon chimpanzee observed
Nigeria-Cameroon chimpanzee observed before 1983
Central chimpanzee
Niaena-Cameroon chimpanzee
Lake
Chad
Estimated range
after
--.
''"^
1983
}^~ it
-^P&BR^^s
Ol
Maroua
<f /
NIGERIA
r^
Gafousr
CHAD
a-.sî
Benouef^S
\
^^^
^'^rpr-^
A
NP & br\
j(Faro NPJ)
V"
Ngaounderel
Fungom FR
Mount Kilum-liim
Community Forest
y'-;
...
O
6N
Ejagham FR-r
Hills
NP
FR-
BAKOSSI
^'^
C
iS-'-T-v
et
WS
DjeremNPo
o
-^
NP (Proposed)
MOUNTAINS
<J'^^Mbam
Bafoussam
Banyang Mbo
Mount Manengouba
Q^ j:
CENTRAL
AFRICAN REPUBLIC
O.
T
Bamenda
Korup
CAMEROON
^-.,
<
Takamanda
FR
Rumpi
Tchabal Mbabo
NP (Proposed)
^-^
r-.'it.j*
ni
,4/
j^-
Bertqua*
Mount Kup6 lER
.v-r--t-,-^
.1--T-
Yaounde
I
i
BIOKO
EQUATORIAL
GUINEA
Douala-'\/r7>.
Edâ WR
V_}
Dja
WR (FaR),
BR & WHS
Cull ul Guinea
^.
r"
CampoNP
"" f
Ebolowa
Ma'a n
MengamS
Gorilla
nation\^le
Boucles du Ntem
FR
ha
/EQUATORIAL,
cPUiNEA
/
8E
306
,^t:
AFRICA: Cameroon
known
some 205-250 weaned Cross
be
to
goriUas, of which about
suspected that the
50
It
is
total population size is slightly
and further surveys are underway. More
larger,
details can be found
There
in
Chapter 7
volume.
of this
the Ebo forest
in
proposed national parki, which appears

of extinction."
population
unknown; measurements
is
to
The taxonomic status
be
of
DNAI,
may
typically yield
was
half the country's land area,
2000, with an estimated decrease of
in
km' per year" Large areas have been
2 220
to shifting cultivation, plantation
lost
development, and
degraded by fuelwood collection
and commercial selective logging.
the southern
In
province of Cameroon, for example, the cumulative
in
number
of this
of the only
Genetic analysis of fresh fecal samples, or freshly
shed hairs (though these
forested
la
male, give ambiguous results.
skull available, a
238 580 km', or
forest fires, or else
also a small population of gorillas
is
north of the Sanaga River
danger
River
Cameroon."
live in
these Ebo gorillas."
some companies have been shown to

illegally over much larger areas than
addition,
trees
granted
in
opens up access
and
hunters
for
are
of forest areas,
who use
arms,'" and increases local
fire
In
fell
their concessions.
Logging causes fragmentation
lower levels
help to resolve the taxonomic status of
logging concessions from 1959 to 1999
of
covered 76 percent of the total forest area."'^^
wire snares
demand
for
bushmeat. Logging routes also serve as conduits
Nigeria-Cameroon cliimpanzees
tor the
The Nigeria-Cameroon chimpanzee occurs
same
forests as the Cross River gorilla, and also has
Cameroon."
a wider distribution within southwest

In
the
in
addition
areas mentioned above,
the
to
Mbo
range includes Korup NP, Banyang

Sanctuary,
proposed
Wildlife
Rumpi
Hills
Wildlife
the north
of
into
income
market towns."
to
urban people, being cheaper than beef
many
threats for
Cameroon's reserves.'" There
of
numbers
of
(proposed), and
Takamanda Forest
Nigeria-Cameroon chimpanzees
some areas
Reserve.'"'
to
" "
in
in
Cameroon. The
populations are thought to be at Korup,
hunting.^'
Chimpanzees were wiped
Kilum-ljim forest
1998.
It
in
killed
Dja Wildlife Reserve. Dja's wildlife
Other large-scale development
the
same geographical range as
western lowland
gorilla,
forests. In 2001, there
35 000 chimpanzees
increases hunting
market
for
intensity
in
this
forest areas
and the
in total in
to
be about
Table 16.2 Central chimpanzee population density estimates
Weaned
Campo-Ma'an NP, Lac Lobeke NP,
Cameroon
Wildlife Reserve, Nki
NP, and
* Their population
km^
1081-1 771"
0.7 10.6-0,91"
0.8 10.6-1.01"
Dja
Douala-Edea Wildlife Reserve,
Gorilla Sanctuary."
in
individuals per
1.2
Protected areas containing central chimpanzees
Mengame
of
the country, which,
Dja Wildlife Reserve
Reserve,
local
the southern moist
i.e.
were estimated
Reserve
Boumba-Bek NP and
the
based on the numbers above, would imply that
Wildlife
is
bushmeat. During the construction
there are at least 30 000 central chimpanzees.*
include
the
in
also under
pressure from logging, with active concessions sur-
also
Cameroon
is
the
1987-
gorillas^'
annually
rounding the reserve, although technically
Central chimpanzees
in
illegal
in
in
has been estimated that about kk
and over 50 chimpanzees^ are
protected buffer zone^''""
Central chimpanzees are distributed
out
northwest Cameroon
Takamanda, and Ebo-Ndokbou.
much
many
in
areas.* Hunting and encroachment are recognized
have declined significantly as a result
Brazzaville estimated that there are around 3 380
over
hunters and an
affordable source of animal protein to rural and
Bakossi Mountains
Population data are limited, but a 2005 workshop
largest
be transported
are indications that Cross River gorilla
Reserve (proposed). Mount Kupe and Mount
Manengouba NP
to
offers both an
Reserve lalso a
Forest
wildlife reserve), the
its
meat
Bushmeat
Outside northern periphery

Dja
mangrove
Lac Lobeke
of Dja
1,1"
Reserve
0.61'"
forest
NP
o.u"
0.17'
densities appear to vary with habitat type, and are
Mengame
Gorilla
0.18'"'*
Sanctuary
particularly high in the Dja area Isee Table 16.2).

0.64^'
Ntibonkeuh
Campo-Ma'an NP
THREATS
The
principal
Cameroon
threats
to
the
great
apes
Boumba-Bek NP and
0.63-0.78^'
Wildlife Reserve. Nki
NP
0.3 IG.2-G.4I'
of
are hunting and logging. ^^ Approximately
307
World Atlas
Map
Cameroon
in
Data sources are provided al the end of
1pE
1>e--.
this
country profile
16E
X
Lake
IN
Species
Cross River
gorilla
id
observed 1989-2004
Cross River gonlla presence alleged
Western lowland
gorilla
Unidentified gorilla species
/:
waz;
Confirmed range
NP & BR
Western lowland
gorilla
laroCia
Estimated range
ir-'--./-;--,
Western lowland
-*T
gorilla
-~-tt^-i;->i jSi^
^^^
Qaroua
CHAD
NIGERIA
BenoueV
NP&B^
:/FaNP^
Bouba
Ndjida
NP
B'N
Ngaoundere
;^
Kagwene Mountain
Tohabal Mbabo
NP (Proposed)
Mbambe
r\ pp
1^
Cross River Gorilla

Sanctuary
"ÎJ
CAMEROON
/ Mount Kilum-ljim
Takamanda
'
Community Forest
QMbam
Bamenda
Nta AN
batouri
Djerem
CENTRAL
6"N
et
AFRICAN REPUBLIC
NP
FR
Bafoussam
Korup
m-
Rump
Hills FR
NANGA
EBOKO
'^^
Bertoua
(Proposed)
BAKASSI
<5-,<i^
PENINSULA ^g7e,,
-4^
^"
DOUMEDIMAKO
Do.a/a
MOUNT
CAMEROON
I
yaoundt
BlOfCO
Douala-Ed6a\
EQUATORIAL
GUINEA
C
(/
//
WR
Mengam4
(T
Sî^
Gonlla Sanctuary
Trinationale
"^^^
EQUATORIAL
GUINEA
lO'E
308
Etmlowa
NP
iNr
of Guinea
Boucles
clesduNtemFR
2"N
Campo-Ma'an
GABON "^
"
!
12E
CONGO
/
14E
16"E
de
-e la Sangha
ar Lnhiko
(Lac
LobSKe MD\
NP)
(I
"
2''N
AFRICA: Cameroon
the
Cameroon-Chad pipeline in 2001-2003. for

oil company Esso reported on several
example, the
cases
it
of
bushmeat purchase by
discovered
in
workers, which
its
the course of attempting to enforce
The net
result of all these factors
among
is
revenues from wildlife-based income
way
in
tax
a similar
Revenue Enhancement
to the existing Forest
in
Protected areas
Formally established protected areas that contain
great ape populations
LEGISLATION
AND CONSERVATION ACTION
Cameroon
in
include the
following national parks, forest reserves, and wild-
[,fe
'"'"'
reserves.
No. 94/01 11994) sets out the country's forestry,
wildlife,
and fishery regulations, and
and chimpanzees as Category

fully
revenue
an inferred
great ape populations
Cameroon.
Law
for a wildlife
Program.'
stronger environmental standards.'"^'
steady decline
The Ministry has plans
enhancement program, which would secure
protected against hunting, capture, or sale,
whole or
in part.
parks and
Nigeria-Cameroon chimpanzees and Cross
gorillas
lists
species, which are
River gorillas:
Protected areas such as national
under the auspices
may be
reserves
wildlife
of the Direction
des Aires Protegees (DFAPI
the
of
la
Faune
status of this area],
is
Djerem
also
responsible for the protection of the country's

biodiversity in general. Article 7 of the
Normes
Document des
calls for a protected buffer
each protected area,
to shield
other activities that might

biodiversity.
it
forest health
Forest Reserve.
Korup
NP,
Mbam
Ejagham
NP,
Reserve,
Banyang Mbo
Wildlife
Fungom
Forest Reserve,
Rumpi
et
Forest
Sanctuary,
Hills Forest
Reserve, and others.
Central chimpanzees and western lowland
zone around
from hunting and
damage
Mone
Nigeria-Cameroon chimpanzees:
et
Ministry of
Environment and Forestry (MINEFI, which
Ithe
recently proposed increasing the protection
established
de
Takamanda Forest Reserve
Environment and Forestry has
Ministry of
in
Lac
gorillas:
Lobeke
Campo-Ma'an
NP,
NP, Dja Wildlife Reserve/Biosphere Reserve,
and
Taxes and permit requirements have
Mengame
NP,
Nki
Sanctuary, and
Gorilla
also been imposed on hunters with the aim of re-
Boumba-Bek NP and
ducing indiscriminate hunting
Central chimpanzees: Douala-Edea Wildlife
Many
in
protected areas."
of the country's forest reserves
were set
Wildlife Reserve.
Reserve.
up by the British colonial administration. The aim
was
to
protect watersheds,
expansion, and
to
restrict
agricultural
conserve areas for future timber
exploitation. Designation as a forest reserve
does not
therefore automatically offer protection from future
logging concessions. Article
Normes
prohibits forest
1 1
of the
Document des
development
activities in
forest reserves as well as conservation areas

their buffer zones,"
but the designation
removed and the area auctioned

The same
236
kml
Several reserves
national boundaries, which
boundary
legislation provides for six types of
some
community forests."" Companies
the largest
close to inter-
means
that territorial
influence conservation outcomes.
the
is
fall
conflicts with neighboring countries can
An example
is
Nigeria-Cameroon boundary disagreement
over the Bakassi Peninsula. This has hampered cooperation between the two countries
the several protected areas
for timber''
logging license to be issued, together with

provision for
and
may be
Of these areas, Dja Wildlife Reserve

at 6
River NP)
in
(e.g.
in
managing
Korup NP, Cross
the transfrontier range of the Cross
River gorilla and Nigeria-Cameroon chimpanzee.
that
break the terms of these licenses can be fined
International support
and disqualified from bidding for further licenses.
The government
Under the supervision
the Ministry of Environ-
cant support from the international conservation
Agence Nationale d'Appui au
community, both nongovernmental and govern-
ment and
Forestry, the
of
Developpement Forestier lANAFOR)

for forest inventory
is
responsible
and management, promotion
of
the use of timber species, soil protection, desertifi-
Cameroon has
received signifi-
mental, for the protection of great apes and the

biodiversity of their habitats.
Some
of the highlights
are listed here.
and forest regeneration." Resources
cation control,
with which
to
mentation
Law
of
of
monitor and enforce the impleNo. 94/01 are, however, scarce.
The Korup Forest Project
NP began
in
in
the mid-1980s.
and around Korup

It
was among
the
309
World Atlas
orate General for International Cooperation

(DGISl and the Foundation for Environment
and Development
(CAMRAILl
in
Cameroon (FEDECI.
in
also partners the
Cameroon
program
WCS
Rail
Company
to control
the illegal
transportation of bushmeat.
government
the
1999,
In
announced the launch

help finance
of Cameroon
new trust fund to
management of
of a
the effective
protected forest areas, and an
US$500 000 was made
of
lake
in
lead projects
Mountains, Cameroon.
policy,
Bethan Morgan
first
major integrated conservation and deve-
lopment projects invested
government
partnership with nongovern-
in
mental organizations INGOsI

Earthlife Foundation,
and then
WWF-The
A series of
later
such investments has been made
Korup
in
NP
and the neighboring Cross River

Division! in Nigeria by the
the
(initially
Global Conservation Organization).
NP
(Oban
UK Department
Development,
International
UK
by the
in
for
European
the
Commission, and others.
ILAGAl,
was formed
1990s
the early
in
to
monitor, track down, and ensure prosecution

of
poachers.^'
In
the
2001,
recorded
first
person was imprisoned after having been

caught trying
to sell a
baby chimpanzee."
of
additional
mammals
in
Cameroon
since
has also assisted Cameroon with
It
gazetting
several
new
protected areas
in
Bakossiland, including the proposed Bakossi
Mountains Wildlife Reserve.'^

In
2002, the Jane Goodall Institute UGI] agreed
to establish a
and
research program
wildlife
(1
150
Mengame
on
located
km'l,
in
Gorilla
the then
Sanctuary
border
the
of
Cameroon and Gabon."

Within and outside the protected areas, the
UK-based Global Witness
is
working with the
Ministry of Environment and Forestry as an
independent observer
improve gov-
help
to
ernance and transparency
and Rational Use
of
of the Zoolo-
San Diego has been conducting
surveys on large
2000.
research,
Conservation and Research
The European Union program. Conservation
in
the forest sector,
Forest Ecosystems
in
concentrating on the issue of illegal
has been active
in
Collaborative agreements have been signed
Central Africa lECOFACI,
logging.''^
the Dja Wildlife Reserve." For example, global
between Cameroon and Birdlife International
positioning systems IGPSI
and between Cameroon and the US-based
and
map
hunters
The
in
were used
paths, snares, and
to locate
camps used
by
the reserve.''"
in
Cameroon
(WCSl has
since 1998, inte-
grating field research, education, and outreach,
as well as support and capacity building of
government
local
World Resources
agencies.'''
Within the
Cameroon-Nigeria border region,
WCS
sup-
UK-based
(FFII
have
all
Mbo
Mbam
et
Djerem
NP and
Living
Earth,
Bristol
Zoo
sponsored research or education
The US-based Bushmeat Project works
in
in
at
of forest
activities."
servation, together with biological surveys

is
aimed
Gardens, and Fauna and Flora International
involve hunters
both countries. Technical support
(WRIj,
and monitoring
resources and biodiversity.
The
ports Cross River gorilla research and con-
to the
Institute
effective conservation
Wildlife Conservation Society
been working
in
forest
to
and fauna protection
eastern Cameroon."
provided
the Banyang
Wildlife Sanctuary, with financing
from
multiple donors including the Dutch Direct-
310
gical Society of
The
WWF
projects.
Endangered Species (CRES)
newly declared
The Israel-based NGO. The Last Great Ape
has
and education
The department
for
of
various protected areas, while
in
WWF-Cameroon
the Bakossi
WWF"
US branches
UK, International, and
Lake Berne, a crater
donation
initial
by
The government
of
Cameroon has
also entered into
agreements with the governments
of
neighboring
and nearby countries, including the following:
AFRICA: Cameroon
Cameroon have established an
Nigeria and
agreement
the Cross River gorilla.
to protect
has also been
partnership
collaborative
include the Durrell Wildlife Conservation Trust's

investigation into
formed between the relevant government

departments, WCS, Fauna and Flora Inter-
German overseas development
national, the
bushmeat supply and demand
Public education and awareness
is
theme
projects and a major
Foundation INCFI.
biodiversity,
Cameroon, CAR, Congo, Chad, Equatorial
reasons - apes and other
Guinea, and Gabon
in
some
traditional
Declaration (see Chapter UJ.
new cross-border
to create
areas
in
outlines plans
It
protected forest
Congo Basin.
the
The Conference
of Central African Moist Forest
Ecosystems (CEFDHACI, coordinated by lUCN-
The World Conservation Union

later
nominated
in
Yaounde, was
an intergovernmental
"
to lead
process based on the Yaounde Declaration.'
The
initiative
the endorsement of the existing 7 300
km^
areas be-
protected
of
tween Cameroon, CAR, and Congo as the Tnnationale de

is
in
are important
The enforcement
of
for
example, led
hunting from
in
Maan community
the
an end
effort to put
to
significant
mid-199^." Similarly,
now making an
are
poaching, and within the
to
Cross River gorilla range, local communities have
agreed
to
ape
protect
populations."
Formal
awareness projects include:
includes:
network
trinational
mammals
hunting ban to protect the sacred
Mount Kupe,
chiefs
protect
both for conservation and symbolic
traditional ceremonies.
decrease
NGOs.
of the national
Local communities are encouraged to
signed the 1999 Yaounde
compo-
nent of most current conservation and development
agency GTZ, and the Nigerian Conservation
all
in
the Cross River gorilla region.
la
Sangha:" the Cameroon sector
Lac Lobeke NP, which supports western
In
Defense
built
of
Animals - Africa IIDA-Africal has
an educational center and
launching a
is
national radio campaign.
The Cross River
Gorilla Project, led by
incorporates a
local
WCS,
education component
lowland gorillas and central chimpanzees;"
consisting of slide shows, posters, and leaflets
focused on the conservation
in
new
the creation of two
national parks
Cameroon: Campo Maan and
Djerem; these were established
pipeline
et
compensa-
offsite
schools and universities.^'
African World Heritage Forest
The Great Apes Project
initiatives
with
Initiative,
the
and the Congo Basin
Project,
groups
of
Forest Partnership. Protected areas involved
Mengame, and Campo Ma'an."
working
in
capacity
is
a loose
mainly
located
building
higher education
in
in
Grands Singes)
(Projet
the Dja area has been involved
in
by
European and Cameroonian students, who

assist
to
in
training
and research
in
topics related
ape protection,
The Bushmeat Project

has provided material
Research and education
Cross River
wildlife.^"
Nature
la
scheme, through an
Cameroonian involvement include the Central
include Dja,
The Club des Amis de

association
Other emerging transfrontier
of the
and other endangered
Chad-
environmental enhancement program.'^
Congo Basin
gorilla
of the
impacts
tion for the biodiversity
Cameroon
in
Mbam
in
for
south Cameroon
use
in
schools and
training workshops.
Various educational and research institutes exist

in
Cameroon.
of
Wildlife
In
particular, the
School
for Training
Garoua, supervised by
Specialists at
the Ministry of Environment and Forestry,

principal school of
The country has

the
its
type
in
the
Francophone Africa."
six universities, of
University of
is
which
at least
Dschang includes research
related to wildlife conservation within
program. The Research
is
forestry
Institute for Agriculture
and Development (IRAD, formerly the

Zootechnical Research)
its
based
Yaounde, under the Ministry
in
Institute for
Nkolbisson near
of Scientific
Research.
Research projects carried out by foreign nationals
The Sanaga-Yong Chimpanzee Rescue Center,

run by
in
In
Defense
August 1999
originally
of
in
dedicated
habilitation of adult
Animals -
Africa,
Central province.
to
the
opened
It
was
rescue and re-
chimpanzees rather than
younger orphans, but young chimpanzees

have more recently been accepted.
chimpanzees
in
The Cameroon
It
held 39
September 200A."
Wildlife
works with the Ministry
Aid
of
Fund (CWAFI
Environment and
Forestry to care for the animals at Yaounde
311
World Atlas
habitat.
As mentioned
Nigeria, this
would
in
the country profile for
establishment
entail the
Okwangwo complex,
As part
the
the
particular by upgrading the
in
protection status of the
Takamanda
development
of the
of a
Takamanda-Mone-Mbulu
(vtinistry
of a
Takamanda-
transfrontier protected area for the
Forest Reserve.
land-use plan for
area, the
WCS
and
Environment and Forestry are
of
currently working on the creation of a protected
area on
Kagwene Mountain,
in
the forests of Mbulu
and Njikwa. Further recommendations include the

urgent need
strengthen
to
enforcement measures
and law
protection
for all
Cross River
gorilla
populations.
Three areas have been identified by the

Ministry as priorities for protection
drill at
the Limbe
in
the Congolian
lowland evergreen forest of the extreme southeast:
Boumba-Bek NP and
Wildlife Centre.
Zoo
at
Mvog-Betsi, and their involvement has
innprovements
led to great
and education
in
Foundation.
Gorilla
existing
illegal logging.
The National Great Ape Survival Plan (NGASP)
Workshop held
in Cameroon suggested that Ebo,

Makombe, Mbulu, Mbargue, Kupe, and Bakossi
forests
were
priorities for great
ape conservation.'"
This sanctuary cares for and rehabilitates
Protection forests are also proposed
bushmeat orphans. Animals are released
mountain region near Mount Cameroon: Etinde,
The UK's
into a restricted area.
Gardens also provides support
The
Limbe
Wildlife
In
January 2004,
it
Bristol
to
Centre
rehabilitating captured great
species.
in
favor of the proposition that an integrated ecological
Bakossiland areas.
apes and other
was
subspecies),
11
one Cross River
western lowland
gorilla.^"
It
In
reserve be declared
2002, the Kupe chiefs voted
in
the
Mount Kupe
in
area.^
caring for
gorillas,
and
receives financial
support from the Arcus Foundation.'
NGO
the western
Mabeta-Moliwe, Kilum Mountain (Mount Oku), and
active
is
in
Zoo
CWAF.
36 chimpanzees (mostly Nigeria-Cameroon
Logging concessions
It
has been recommended that logging companies
be requested
to
agement plans
produce great-ape-sensitive manfor their concessions. This
would
Pandrillus organized the
include controlling illegal hunting and financing law
repatriation of two confiscated western low-
enforcement, as well as providing protein alter-
In
2004, the
land gorillas from Nigeria to

first
Cameroon
- the
time the two countries had cooperated to
resolve
an
trade
a species listed by the Convention on
in
natives to workers and local
in
Endangered Species
of
Wild Fauna and Flora (CITES)."
Capacity building
For Cross River gorillas,
that a
it
government departments,
In
Following the Second International Worl<shop and
lack of
the Korup
governments
of
ledged the need
Cameroon and
to
protect
in
2003,"^'" the
Nigeria acl^now-
Cross River gorilla
universities,
is
built in
and NGOs."
general, reserves and parks sufferfrom a chronic
Conference on the Conservation of the Cross River

Limbe, Cameroon
established, and
and conservation capacity
Protected areas
in
has been recommended
management committee be
that research
Gorilla held
communities affected
by logging.'"
instance of illegal cross-border
International Trade
312
of
protected area boundaries might help to discourage
works
2001 with the sup-
in
US-based
the
of
also
where the Michael Leo Rion
Sanctuary was opened

port
CWAF
at the zoo.
the Mefou NP,
animal welfare
in
Lobeke
Wildlife Reserve, Lac
NP, and Nki NP. Better demarcation
staff,
equipment, and infrastructure. Only
NP
has both
chief warden. Action
is
management
needed
to
plan and
strengthen the
Ministry of Environment and Forestry's ability to
enforce
forestry
legislation,
including
hunting
AFRICA: Cameroon
laws.'"''
berand
'"'^^
In
particular, an increase in the
num-
Education and community development

information and education campaigns are needed
training of ministry wildlife monitoring staff
has been widely recommended."'"
on a
large
scale
them about
Research
to
Further, better-coordinated research and monitor-
Training
workshops
groups
should
needed on ape distributions and populations
ing is
throughout Cameroon.'"^*
search
and population
into the ecology, distribution,
expanded.^'
the south
underway."
is
study of the Ebo gorilla population
wildlife
for
also
sources as alternatives
Cross River gorillas should be
biology of the
tell
Cameroonians
the
of
of
great apes and
protection
laws.
community members and

organized,
be
to
cover
conservation issues and the use of other protein
particular, basic re-
In
inform
to
endangered status
the
should be given
engaged
in
bushmeat."'
to
'
Support
developing alternative methods
income
generating
of
to
to
hunting.
for
people
currently
'*
FURTHER READING
Bikie, H.,
Collomb, J-G., Djomo,
L.,
Minnemeyer,
S.,
Ngoufo,
R.,
Nguiffo, S. 120001
An Overview
of Logging in
Cameroon. Global Forest Watch, http://www.globalforestwatch.org/common/cameroon/english/report.pdf.

Accessed June 12 2005.
Comiskey,
J.A.,
Sunderland, T.C.H., Sunderland-Groves,
J.L.,
eds 120031 Takamanda: The Biodiversity of an African
Rainforest. Smithsonian Institution, Washington, DC.
Dupain,
J.,
Guislain,
densities
in
P.,
Nguenang, G.M., De Vleeschouwer,
K.,
Van Elsacker,
(2004) High
a non-protected area on the northern periphery of the Dja Faunal Reserve,
chimpanzee and
gorilla
Cameroon. Oryx 38
121:
209-216.
Global Witness 120051 Forest
December 2003-June
Law Enforcement
in
Cameroon: Third Summary Report of the Independent Observer
2005. http://wwM(.globalwitne55.org/reports/show.php/en. 00072. html. Accessed June 12
2005.
Gonder, M.K., Dates,
J.F.,
Disotell, T.R., Forstner, M.R., Morales, J.C, Melnick, D.J. 119971
A new West
African
chimpanzee subspecies? Wature 388: 337.

Matthews,
A.,
Matthews, A. 120041 Sun/ey
troglodytes]
MINEF
12003)
in
of gorillas {Gorilla gorilla gorilla]
southwestern Cameroon. Primates
Cameroon Action Plan
Proceedings
J.L., Jaff,
of the
B.,
of the
and chimpanzees IPan troglodytes
15-24.
for the Survival of Great Apes
Cameroon, March 18-20. Cameroon Ministry

Sunderland-Groves,
it5:
and Endangered Primates. Workshop
report, Mfou,
Environment and Forestry IMINEF).
eds 12004) Developing a Conservation Strategy for the Cross River
2nd International Workshop and Conference on the Cross River
Gorilla. Wildlife
Gorilla.
Conservation
Society.
MAP DATA SOURCES

Maps
16.3a and b Great apes data are based on the following source:
Butynski.T.M. 12001) Africa's great apes.

Stevens,
E.F.,
Arluke,
A., eds.
In:
T.S.,
Hutchms,
M.,
Maple,
T.L.,
Norton, B, Rowan,
A.,
Great Apes and Humans: The Ethics of Coexistence. SmUhsonian Institution Press,

With additional data by personal communication from Bergl, R. and Sunderland-Groves,
J L. 120051
and from the
following sources:
Dowsett-Lemaire,
in
Haltord,
et
F.,
Dowsett, R.J. 12001) A
new
western Cameroon. African Primates

T.,
Ekodeck,
H.,
Sock,
B.,
Dame,
5:
population of gorillas Gorilla gorilla and other endangered primates

3-7.
M., Auzel,
P.
12003) Statut des populations de gorilles /Gorilla gorilla gorilla/
de chimpanzes IPan troglodytes troglodytes/ dans
Cameroun:
densite,
distribution,
le
Sanctuaire a Gorilles de Mengame, Province du Sud,
pressions et conservation.
MINEF and Jane
Goodall Institute, Yaounde.
http://www.janegoodall.net/news/assets/RapportGrandsingesMengamefinal2003.pdf. Accessed August
2004.
Morgan,
B.J. 12004)
The
gorillas of the
Ebo
forest,
engli5h/gjournal/texte/28ebo.html. Accessed
Cameroon.
Gorilla Journal 28: 12-14. http://www.berggorilla.de/
November 24
2004.
313
World Atlas
Sunderland-Groves,
in
J.L.,
Maisels,
Ekinde, A. 120031 Surveys of the Cross River gorilla and chimpanzee populations
F.,
Takamanda Forest Reserve, Cameroon.
Comiskey,
In:
Takamanda: The Biodiversity of an African
J. A.,
Sunderland, T.C.H., Sunderland-Groves,
Rainforest- Smithsonian Institution, Washington. DC. pp.
J.L.,
eds,
129-UO.
http://nationalzoo.si.edu/ConservationAndSc1ence/MAB/re5earchprojects/appliedconservat10n/we5tafrica/
Takamandabook/Chapter_9.pdf. Accessed July 13 2004.

Various authors (2003) Draft
map
GRASP
forw/orking purposes prepared at
meeting, Cameroon, March 18-20 2003.
ACKNOWLEDGMENTS
Many thanks
Jean LaGarde
to
Betti (University of
DschangI, John Ngong Fonweban (University of DschangI, Roger
Fotso (Wildlife Conservation Society), Elizabeth Gadsby (Pandrillus), Bethan Morgan [Zoological Society of San Diego),
John F Dates (Hunter College,

Society) for their valuable
New York),
City University of
comments on
and Jacqueline Sunderland-Groves
the draft of this section.
Thanks also
[Wildlife
Conservation
Barry [Tropical Biology
to Brigid
Association) for editorial assistance.
AUTHORS
Patrice Taah Ngalla,
Limbe Botanical and Zoological Gardens

Caldecott, UNEP World Conservation Monitoring Centre
Lera Miles.
Julian
CENTRAL AFRICAN REPUBLIC

Nigel Varpi-
the world, with a
The Central African Republic [CAR]
is
a landlocked
country, covering 622 984 km'. Barthelemy
Boganda
was
killed in
led the country to independence, but
1959 shortly before secession from France.
became
David Dacko
dent
of
the
first
1960,
In
independent presi-
CAR. Elections were held
in
1964,
in
which
Dacko was the only candidate, representing the only

party." Deteriorating
to a
economic circumstances
led
1966 coup d'etat led by Colonel Jean-Bedel
Bokassa,
who
power;
1972 he declared himself president for
life.
In
in
later
1977, Bokassa
renamed
obtained virtually absolute
was crowned as emperor
of
country, the Central African Empire.
Opposition
to
massacres
in
his
erratic
rule
led
to
riots
and
1979, followed by a French-backed
coup that reinstalled Dacko as president and
pectancy
army
mutinies, foreign interventions, and
Index ranked
and a
life
ex-
40 years." The economic

recent years due to
in
severe political disturbances, combined with violence and
looting.
product (GDP)
In
the gross domestic
2002,
was U5$1.1
national income
billion
and the gross
(GNU per person was
US$300. " The major sources
less than
of national
are mining [mostly for diamonds)
in
income
the southwest,
center, and north of the country, and the sale of
timber from logging, mainly
in
the moist forests of
the southwest. The population
be about 3.8 million

1.6 percent.'^
in
Most
in
was estimated
to
2001,^^ with a growth rate of
the people are concentrated
of
the cities and rural areas of the south and west,
leaving the north
and east largely unpopulated.
The Aka people

Bayaka)
(also known as Ba'Aka and

number about 20 000 in CAR and inhabit
coups d'etat occurred periodically during the 1980s,
the southwestern rain forests and savannas. Like
1990s, and into the 2000s, as the country struggled
the other forest-dwelling hunter-gatherer peoples
to
develop democratic institutions; the current head
of state, Francois Bozize
through a coup
in
himself
came
2003. The constitution
to
power
was then
suspended, but following a general election held
in
was returned as president.

among the least developed countries
in
early 2005 Bozize
CAR
3U
at birth of only
2004,*^
in
situation has deteriorated
attowed the republic to be restored. Popular unrest,

strikes,
Human Development
169th out of 177 countries
is
of Central Africa (the
the
BaKa
Twa and the Mbuti
of the
Congo [DRC],
of
of the
all of
Cameroon and Gabon,

Democratic Republic
whom
are sometimes
called 'Pygmies'), they are often subordinate to their
Bantu neighbors. Their homelands are now being

logged, hunted, and settled by outsiders, leading to
AFRICA: Central African Republic
an erosion
of their Livelihoods
including
the
loss
and cultural
Integrity,
hunting
traditional
of
skills,
techniques, and technologies. The latter include
animals such as duiker
driving
and
nets,
into
hunting with spear and bow-and-arrow.
about 229 000 km'
forest,
more than
were estimated
cover
to
2000, or 32 percent of the land
in
CAR
area." Most of
too dry for closed canopy rain
is
which only ever occurred
a narrow band
in
along the southern edge of the country, amounting

to
about 8 percent
area."
of the land
It
is
now
and the capital
Bangui, and the southeastern
city
as
undetermined
yet
subspecies
of
Nabolongo.' "
Bangassou
'
forest
forms part
The island
the
in
southern
of the
There no longer seem
to
be any
Bangassou
in
and the island population continues
itself."
percent of land use
in
concentrated
the
in
only accounts for about 3
CAR
Based on
mean
a 1985 map.^
be
density of 0.25 individuals per
may be
was estimated
it
map
vegetation
through different
of transects
1984-1985. extrapolated over a 1967
in
modified to take account of later
human
The western lowland
between 4 806 and 7 830
gorilla [Gorilla gorilla gorilla]
of forest
that 9 000 gorillas
present over 36 000 kml^' Based on nest
counts over 783 km'

habitat types
overall.''
and the central and eastern subspecies
to
Numbers
in
is
forest
hunted by Congolese."
patchy and degraded.' and Isolated from other rain
It
south of
to the
is
square kilometer occurring over the area
south, even though
an
Mboumou River, which

border of CAR with DRC.
Bangassou region." The Bangassou forests are

forests by farmland, which
of
(presumably
res-
southwest corner, between Gamboula
tricted to the
poorly
Is
known and there are few recent records.

A population of about 40 chimpanzees
eastern) has also been reported from the island
Forests and woodland savannas of

10 percent canopy cover
the country, but their current distribution
habitation,
km'
it
was estimated
that there
were
gorillas in the southern-
of
the
most
chimpanzee [Pan troglodytes troglodytes and
P.t.
positive picture than that given by the 1980 estimate
schweinfurthii] are
is
found
all
in
CAR. The country
of
northeastern edge of the range of the
at the
western lowland
Neither gorillas
censused
at a
and central chimpanzee.
gorilla
nor chimpanzees
have
but the
national level."
been
Dzanga-
6 000
500 gorillas,"
An estimate published
population
1987 gives a national
in
chimpanzees
of
based on the area
800-1 000. again
at
of suitable habitat." In a
review, however, the estimate
was 800-1 000
Sangha area has been surveyed more thoroughly
central subspecies alone, as well as an
than most other sites
number
CAR.'
in
They
restricted to the forests of the southwest.
occur
both the Dzanga and Ndoki sectors of
in
Dzanga-Ndoki National Park (NPI

the
(3
are
Gorillas
11
222 km'l.
'
declared an
and
In
the Ngotto forest, which
reserve
integral
in
1996 and
unknown
subspecies." The difference
for the eastern
both cases reflects an improvement
rather than an actual increase
in
knowledge
numbers.
in
THREATS
was
An estimated 300 km'
the total] are cleared
is
2003
for the
in
Dzanga-Sangha Dense Forest Special Reserve
359 km'l.^
in
more
of forest alone." This is a
of forest (about 0.1

in
CAR each
percent of
year." Logging
proposed national park.'" and there are several
concessions cover 50-75 percent
records of gorillas from the surrounding area.
forests.^'in the rain-forested southwest. 86 percent
The central chimpanzee also occurs
in
the
forested southwest, but not abundantly
so.'''
density
square
of
kilometer
in
only
0.16
individuals
was reported
in
per
the Dzanga-Ndoki
NP
1996-1997,' and 0.44 individuals per square
kilometer
in
the
Ngotto Classified
Forest
in
1998-1999.' Central chimpanzees have also been
reported at one location
eastern chimpanzee
of the
is
in
the northwest." The
reported from the forests
southeast, which have never been properly
surveyed for the species."
one recorded
'
and again, there
locality in the north.
panzees were previously known
in
is
Eastern chimcentral parts of
of forests
had been allocated
2000." High transport costs
CAR tends
of
to
of the
remaining
concessions by
to
mean
that logging in
be highly selective, but the operation
these concessions inevitably opens up previously
inaccessible areas to hunting.
chimpanzees are vulnerable
Both gorillas and

hunting, logging,
to
and forest clearance, so they are assumed

decline, although the rate of loss
There
is little
systematically
known
in
is
to
be
in
unknown.
evidence that gorillas are hunted
CAR.'''
prey, albeit at an
meat trade
is
" but chimpanzees are
unknown
level.
The bush-
considered the greatest threat to
wildlife conservation in
CAR, even inside the pro-
315
World Atlas
Map
in
the Central African Republic
this
country profile
1\
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r
o
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ID
> 1
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316
^]îVP
ISCD
6'^
'P li
i ;^
^
=
rot:--,
Af-
-'^.^s-i
ô
tected area system.
bushmeat
forest areas,
In
is
less expensive than chicken, goat, or kinin caterpillars (an
Aka
important food source for forest-dwelling
Firearms have been cheap
people!.''
most
years, '^ but
in
local hunters in the forest

^'
and cable snares.^'
use nets
Capture and injury by snares
placed for other animals
ous hunting threat
is
probably the most seri-
apes
to great
CAR.^
in
study
Dzanga-Sangha Special Reserve found
the
recent
in
that at
any one time approximately 60 people were using

wire snares
in
an area
of
about
east,
rifles,
where
gangs
more common
is
the north and
in
poachers from Sudan." Hunting pressure
of
was reported
be very high
to
in
the Ngotto forest
in
the 1990s, particularly near the town of Bambio,''
somewhat." The Aka
but to have later decreased
and
Bofi people living in the reserve
appear
focus
to
on small game, and often hunt with nets designed

for this purpose. ''
The Bangassou
hunted throughout and unless

will
become an
'empty'
this
forest
changes soon,
''
forest.'"-
heavily
is
Hunting
in
forest.
Preliminary steps for the
ratification of the
Convention on Wetlands
Zemongo Faunal
Reserve, has
also been intense.
Environment, Waters, Forests,
of the
Fishing
is
responsible for wildlife

in
CAR. These are governed by Ordinance No. 84.0^5

of July
27 198i, which deals with the protection of
and Law No. 90.003
pertains to the
of
June 9 1990, which
Central African
Customary hunting
territory
Forestry Code.
authorized throughout the
is
CAR, with the exception
of
integral
of
reserves and national parks.-" Great apes are listed

in
Category A as 'completely protected' under
forest
agreements
Diversity
International Trade
in
in
Combat
1995,
tlie
in
1980, the
Ndoki
in
is
Cameroon
la
Sangha
Cameroon and
The Dzanga-
(TNSl.
NP
the Nouabale-Ndoki
which together form the 7 300 km' core

conservation area.
the tripartite
NP
of
Congo,
in
of the
TNS
other achievements,
TNS agreement,
2000, has resulted
change
Among
in
known
contiguous with the Lac Lobeke
signed
December
in
ranger patrols and ex-
in joint
of intelligence, leading to
some
successful
antipoaching missions.
Protected areas
There are three principal categories

area
in
CAR:
strict
There
of protected
nature reserves (one
site),
and faunal reserves
[eight
(five sites),
also one special reserve (Dzanga-
is
area system includes almost
private reserve
in
1 1
percent
of
the land
area, only 32 percent of the protected areas are
thought
to
be adequately managed, and three are
completely unmanaged.' The latter include the
Zemongo Faunal
panzee
is
still
Reserve, where the eastern chim-
believed to occur About 50 areas,
covering around
percent of the national territory,
have been gazetted as forest reserves

or
('classified'
gazetted' forests), intended for the sustainable
production of forest products.' "

actively
Few
of
these are
managed.'
The Dzanga-Ndoki NP
in
of the
220 km' and
country covers
the extreme southis
sectors, which are joined to the 3 359
Convention on
Sangha Special Reserve. The topography
UN
of
Wild
Convention
Dzanga sector
patchwork
of
is
relatively flat,
Nature and
were
here
of
1980 (under which one
site
has
been designated, the Manovo-Gounda-St Floris
and
of
the
comprises a
it
forest, with
undergrowth.' Parts of the park
selectively logged prior to 1982.'

is
km' Dzanga-
primary and secondary
1969, and the World Heritage
1996,
divided
Dzanga (495 km'] and Ndoki (725 km')
much herbaceous
in
Convention
MP
the African
Desertification
Natural Resources
Ramsar
national
Convention on
Endangered Species
Fauna and Flora ICITESI

to
first
political
the Ngotto
a three-way transfrontier reserve
as Trinationale de
into the
or acceded to the
ratified
Biological
the
collaborates with Congo and
managing
west
CAR
was proposed as
to
in
Site.'
Ordinance No. 84.045.
International
of Inter-
(Avakaba Presidential Park). While the protected
conservation and the use of natural resources
wildlife,
The Mbaere-Bodingue area
Sangha Special Reserve) and one
Hunting, and
was delayed due
but final ratification

instability.
sites)
The Ministry
Basse-Lobaye
is
national Importance (Ramsar) have been conducted
national parks
LEGISLATION
which
it
the
east and southeast of the country, including the

buffer zone of the
of
near the Ngotto
CAR
often undertaken by well organized
is
it
Reserves under UNESCO's Man and Biosphere
(MABI Programme, one
000 km'."
Hunting with firearms, including automatic

assault
NPI. The country has also designated two Biosphere
"The
climate
characterized by a dry season of three
months (December-February) and

season, with a drier period
in
long
rainy
June-July.' Gorilla
317
World Atlas
chimpanzees. Based on these densities, an
mated population
of
380 weaned chimpanzees occurs
Mbaere-Bodingue NP 1872
Conservation and
logging operation
1989'!
in
field projects
when
Dzanga-Sangha
Republic.
conducted
densities of 1.6 individuals per square kilometer
human
have been recorded within the
2000''
times
corded
to forests within the
and
traditional
is
ac-
national park, while
safari hunting, agroforestry develop-
ment, and commercial logging have occurred

special reserve."'"
was developed
In
in
the
scheme
the early 1990s, a
share revenues from tourism
to
in
the reserve and park, with 40 percent going to local
communities and 50 percent

for
tration
salaries,
to reserve
Since 2001, the reopening
of a
within the reserve has led to a
and the reopening
siders,
adminis-
upkeep, and maintenance.^'
in
poaching,
CAR
of
roads
ivory
chimpanzees are also reported
forest,' part of
may become
Dzanga-Sangha.
swamp
forest
is
It
large
in
dense semi-
limits
enhanced by
its
More than
13 primates) and
found
in
of
the
Sudanian wooded
position astride the three
major
1 1
5 species of
mammal (including
more than 320 species
the Ngotto forest," which
is
of bird are
regarded as an
area of national importance for the conservation of

at least four
monkey
species.'" Estimated gorilla
densities are lower at Ngotto 10.34-0.40

gorillas per km'l than in the
318
a fairly
CAR
nationals.
constant presence
Dzanga-Sangha since the early 1990s, despite
economic and
quent
political fluctuations
instability of the
Environmental education
undertaken
in
and the conse-
developing tourism sector
have also been
activities
the area. Gorilla research,
in
par-
has contributed significant development
participants." Research into
chimpanzee ecology
and behavior has been more limited

study on
feasibility
tourism
in
CAR.' A
in
habituation of apes to eco-
the Ngotto forest
was
carried out
in
and the hunting pressure too high
A number
supported conservation projects

Ndoki
for success."
of international organizations
have
Dzanga-
the
at
NP and Dzanga-Sangha Special Reserve, inWWF-The Global Conservation Organization,
cluding
Bank, the US Agency for International Development
the northern part of
biogeographical zones of Central, West, and East

Africa.
Ape research has had

at
to habituate gorillas
tourism by teams including
the Wildlife Consen/ation Society [WCSl, the World
savanna, the area has high species richness, which

is
has included successful efforts

for
Raphia
of
the Guineo-Congolian forest block, and bordering
southern
at
MBaere and
tracts
along the
Bodingue Rivers. Situated
the
has been taking place
of this
of
1997,
three study areas within the Dzanga-Ndoki NP, and
the
covering about
Ngotto,
NP.'
deciduous forest with

IPalmael
which
the second largest moist forest area
is
after
Remis
(e.g.
2000-2001, but found that densities were too low
Ivlbaere-Bodingue
10 000 km",
Much
"I.
on the gorillas
opportunities for local, national, and international
influx of out-
network
both the special reserve and the
Gorillas and
in
activities
huge
of a
national park.^
from Ngotto
and foraging, and
into nesting, feeding
ticular,
commercial bushmeat hunting and

in
the
in
Research has since been
logging concession
throughout the area. There has been a resultant
upsurge
forest.
in
Fay
(e.g.
behavior of gorillas and the impact
ranging
parl<; that is
CAR began
in
extensive surveys
revealed high densities of animals
the Central African
the density of chimpanzees.^ Full protection
the proposed
in
km'l.'
Studies on western lowland gorillas

the mid-1980s,
esti-
295-350 weaned gorillas and
weaned
Dzanga-Sangha
region,
and the German overseas development
lUSAID],
agency GTZ.
WWF
supports the administration
and development
project. Activities are
management,
protected area
sustainable forestry
tourism,
new
have
wildlife
guards, tourist guides, and trackers, and develop-
ment
of
an antipoaching strategy for the area. There
are also two gorilla habituation sites
Sangha area
Hokou. The
in
the Dzanga-
Mondika Research Station and Bai
latter
WWF tourism
has been developed
vegetation there." The opposite pattern
health monitoring of gorillas.^"
in
management, and
included recruitment and training of
ful
seen
focused on
development,
rural
applied ecological and social research, and
probably due to the lesser abundance of herbaceous

is
of
the complex as part of an integrated conservation
for a
success-
program, which includes extensive
The CAR component

Ecosystems
of
the administration of the Ngotto forest until
drawal
was
in
up a
the government.
triangle
plan and a set of general
An area
forest
of
of
ECOFAC and
872 l<m^ situated within
chimpanzees, has been set aside

is
patrolled by a
and trained by the
team
of
and central
gorillas
for total protection
ecoguards' recruited
Africa should
to
The Bangassou
forest,
where the eastern
chimpanzee has been recorded, was the subject

US$3.5 million community conservation
of
most ape
that
western equatorial
in
be focused on formally protected
where great ape populations are
areas,
be viable
likely
the long run, with an immediate
in
large investment
law enforcement
in
un-
to tackle
poaching and, over the longer term,
controlled
building of national capacity to conduct all aspects
management."
of protected area
One
assessed
study
CAR based on
measures including threats from
variety of
mining,
hunting,
logging,
farming, villages and
grazing,
significance,
biodiversity
and the effectiveness
area,
conservation
the
potential of the protected areas of
roads,
project.'
recommend
authors
conservation investment
between the Bodingue and
M'Baere Rivers that supports
and
with-
sustainable logging operation,
pilot
management
operating conditions established by
its
2003." One of the objectives of the project
to set
based on
Forest
Central Africa (ECOFACI, supported
in
Some
European Union
of the
program, Conservation and Rational Use
of its
integrity
the
of
management. The
Dzanga-Ndoki NP and the Dzanga-Sangha Special
initiative
Reserve had the highest ratings, making them,
financed by the Global Environment Facility IGEF).
with the high density of gorillas they support, a
There
law enforcement
little
is
authorities at present, but
GEF
next phase of the
it
by
the
local
planned that the
is
project will provide for the
recruitment and deployment of paid ecoguards.^"
No sanctuaries
for captive great
apes
"
CAR.'
exist in
priority for continuing great
As
yet there
great apes
ECOFAC
program has recommended
Mbaere-
Bodingue area
in
highlighted
CAR, nor are great apes known
be
number of
authors, however, regarding specific areas

gorillas
to
any national environmental plans.
in
Recommendations have been made by
It
the
that
Ngotto forest be given national
has also been suggested that
logging companies leave an undisturbed strip of
no conservation action plan for the
is
in
to the set of
protected areas under consideration, the
park status.'" "
ape conservation and
research efforts.'^^ Potentially adding
where
forest connecting this area with Dzanga-Ndoki.'"
new
effort to
protect
is to
in
chimpanzees has also been suggested;^^
be hoped that the
achieving
and chimpanzees occur, and regarding
conserve the Bangassou forest
GEF
project
is
A
to
it
instrumental
this.
Increased wildlife law enforcement
is
fre-
the region as a whole for great ape conservation
quently seen as a priority over the country as a
measures.
whole
CAR
will
need continued financial as well as
Blom
(e.g.
e( al. 2D0i').
technical assistance to deal with ape conservation
should be compelled
and protected area management. There
hunting,
is
a real
need for capacity building, regional collaboration,

political
commitment, and the development
sustainable
long-term
funding
of
mechanisms.'
Further information on ape population sizes and

distribution
CAR would
in
greatly
assist
in
transport,
has also been
to control illegal
bushmeat
and consumption on their
concessions. Meanwhile, funding for efforts to control
ated
hunting and the Ebola virus should be generin
developed countries. Regional governments
could then be offered economic Incentives for ape

protection, linking aid and debt relief to verifiable
measures
conservation planning.^^
It
suggested that logging and mining companies
of
conservation performance.'*
FURTHER READING
Blom,
A,,
AlmasI,
A.,
Heitkbnig, I.M.A., ef
al.
12001
A survey
of the
apes
African Republic: a comparison between the census and survey

gorilla]
Blom,
A.,
in
the Dzanga-Ndoki National Park, Central
methods
of estimating the gorilla [Gorilla gorilla
and chimpanzee [Pan troglodytes] nest group density. African Journal of Ecology 2'): 98-105.
Yamlndou,
J.,
Conservation ^^R
Prins, H.H.T. 12004) Status of the protected areas of the Central African Republic. Biological
|4|:
479-487.
Carroll, R.W. (1988) Relative density, range extension,

gorilla] in the
Dzanga-Sangha region
of
and conservation potential
of
the lowland gorilla [Gorilla gorilla
southwestern Central African Republic. Mammalia 52
(31:
309-323.
319
World Atlas
Fay, J.M. (1989) Partial completion of a
Mammalia
53:
census
of
lowland gorilla [GoriUa
Remis, M.J. 11997) Ranging and grouping patterns of a western lowland

Republic.
g. gorilla] in
the Central African Republic.
203-2M.
American Journal of Primatology 3Z:
Remis, M.J. (2000) Preliminary assessment
of the
1 1
gorilla
group
at Bai
Hokou, Central African
1-133.
impacts
of
human
activities
on gorillas Gorilla gorilla gorilla and
other wildlife at Dzanga-Sangha Resen/e, Central African Republic. Oryx3U
111:
56-65.
MAP DATA SOURCES

Map
^b.l^
Great apes data are based on the following source:
Butynski, T.M. 12001) Africa's great apes.

Stevens,
E.F.,
Arluke,
A.,
In:
eds. Great Apes
and Humans: The
T.S.,
Hutchins, M., Maple.
T.L..
Norton,
B..
Rowan,
A.,

With additional data by personal communication from Maisels,
Blom,
A.,
Almasi,
A.,
F.
I200AI and from the following sources:
Heitkbnig, I.M.A., Kpanou, J-B., Prins, H.H.T. 12001)
A sureey
of
the apes
in
the Dzanga-Ndoki
National Park, Central African Republic: a comparison between the census and survey methods of estimating
the gorilla [Gorilla gorilla gorilla] and
chimpanzee [Pan troglodytes] nest group
density. African
Journal of
fco/ogy 39: 98-105,

Brugiere,
D.,
Sakom,
Population density and nesting behaviour of lowland gorillas [Gorilla gorilla gorilla]
D. 12001)
in
the Ngotto Forest, Central African Republic. Journal of Zoology 255: 251-259.
Brugiere,
D.,
Sakom,
Gautier-Hion, A. 12005) The conservation significance of the proposed Mbaere-Bodingue
D.,
National Park, Central African Republic, with special emphasis on
Consen/ation
its
primate community. Biodiversity and
505-522.
121:
ACKNOWLEDGMENTS
Many thanks
to Allard
Blom IWWF-US], David Brugiere ISECA-BRLi Consulting Company], Fiona Maisels
Conservation Society], and Elizabeth A. Williamson (University
of Stirling] for their
valuable
comments on
(Wildlife
the draft of
this section.
AUTHOR
Nigel Varty,
UNEP World
REPUBLIC OF THE CONGO

Nigel Varty

The Republic
Congo
of the
is
cities of Brazzaville,
Central African
country with a land area of 3A2 000 knn^

lation
a
was estimated
growth rate
at 2.95 million in
of 1.5 percent.' Its
product IGDPl for 2002

billion
Its
popu-
at
US$3.0
with a gross nationalincomelGNI] of
US$720
per person." Congo
is
leum producers, with
one
of Africa's
significant
main petro-
potential for
further offshore development. Petroleum (mostly

offshore]
sources
and timber were Congo's two largest

of foreign
exchange
in
2002, while agri-
culture accounted for only 10 percent of GDP.'
than half of the population lives
320
in
More
the southern
of rural
Pointe-Noire, and Loubomo.
Congo has fewer than four
inhabi-
tants per square kilometer.
Congo
2003 with
gross domestic
was estimated
Much
is
one
of the
most densely forested
countries on the African continent. Forests cover
220 600
km^ or about bU.b percent of the land area,

some 830 km' of plantations." In 2003,
including
permanent crops and arable land were estimated

to
account for only 0.6 percent
of the total land
coven' The main forest areas are the
(Maiombe] and ChaiUu massifs
in
Mayombe
the southwest
(which are southern extensions of the Lower Guinea

forest block),
in
and the Sangha and Likouala regions
the north (which are part of the
Congo
Basinl.
AFRICA: Congo
Independence from the French was
when
1960
Republic
Congo. Until 1990. the dominant
of the
system was based on an interpretation
political
of
ratified in
Congo became the
the former Middle
was abandoned, and a more

was elected in 1992.'
outbreak of civil war in 1997, former
Marxism, but
this
market-oriented government
Following the
was
Marxist President Sassou-Nguesso
period
but a
of
civil
unrest followed
elected,
the
until
beginning of this century. Negotiations eventually

led to a
new
draft constitution,
by referendum
tion
won
which was approved
bordering the
chimpanzee release
by Sassou-Nguesso, and subsequently
more
allowing
Most
stable conditions to prevail.
people displaced by the war have returned, mal-
and death rates have dropped, the
nutrition
rail
between Brazzaville and Pointe-Noire has
link
The River Louvandzl,
2002," leading to a national elec-
in
reopened, and the security situation

improving."
signed a
new commitment
government, and
into society
slowly
is
March 2003, the main rebel group
In
peace with the
to
disarmament and reintegration
its
has commenced.
in
BenoTt
Goossens/HELP
clearing and
International
surrounding Marantaceae forests, a
its
study area estimated to support 500 gorillas."

Social stability, high birth rate, and apparent low
have
mortality
infant
was doing
population
in
Congo. The Congo and

of
the range
proportion of the world's western

live in
Congo. They
the densely forested northern region'ând,
forested areas
Congo, and
'^
gorillas,^
'
but
it
is
feared to
have decreased markedly by the early part

21st century. While the 1990 figure
larger than the 1980 estimate of
the
in
been estimated
north, the national population has
000-a 000
in
the southwest of the country.
in
Based on 1990 habitat maps and censuses
is
of the
to
30 000
of
000,^^ the
these gorillas are found
Odzala-Koukoua National Park
known density
found
in
Over a
in
thought
is
aux
of the Likouala
They also occur
km^
the i 190
Nouabale-Ndoki NP, and the surrounding
of
[NPl.
The highest
lowland gorillas (11.3/km')
month period
in
populations
the
forests.'"
be the most
to
is
199i and 1995, A27
southwest Democratic Republic
in
Congo IDRCI are not
numerous
forest
in
block, '^ but these
hunting,^
and
it
Lekoli-Pandaka Faunal Reserve and M'boko Hunting Reserve.
The resulting protected area covers
13^56 km^ and includes the Maya Nord
salt
of the protected
areas
in
which they occur'"

Central chimpanzees occur
it
in
the north and
Congo along the border with Gabon but

widespread in the central region.' " In 1991,
southwest
of
was thought
chimpanzees
that there
were only
000-5 000
Congo," but since then, the
in
mates have been
revised, with a
esti-
2003 figure
around 10 000 chimpanzees.'" The highest

corded density of this species
Chimpanzees are
absorb the adjacent
be
to
has been suggested that their fate
depends on the success
was expanded
to
are considered
vulnerable because of logging and associated
been reported
2001
be
to
Mayombe
the Kouilou Basin part of the
nests were recorded along line transects.^ The park

in
of
extinct after all." In the
have also been reported
gorillas
past,
are less
the
in
the park's extensive Marantaceae forests.

five
'^
River.''
The Oubangui River
substantially
000-3
actual population trend has been negative."
Up
Herbes
easterly limit to the range of the species, unless the
large
to a lesser extent, central parts of
3^
area of north
and both east and west
in
lowland gorillas are thought to
at
swamp
Rivers,
occur
chimpanzee.
in
have also been
gorillas
and central chimpanzee [Pan troglodytes
[Goritla
gorilla
both the western lowland gorilla and central
occur
of
the vast Likouala
Congo between the Oubangui and Sangha
lowland
Oubangui Rivers form the eastern edge
Ebola
of a devastating
central
troglodytes]
of
in
sub-
this
gorilla
western
gorilla]
reported
that
September 2004,
the park Isee below).
High densities
The
indicated
well." By
however, there were fears
epidemic
site
the Conkouati area.
swamp
in
forests,
been recorded
in
in
Central Africa has
Odzala-Koukoua
not
uncommon
and densities
of
of
re-
NP
in
12.2/km'l.'
the Likouala
0.1-1.3/km' have
the Lac Tele/Likouala-aux-Herbes
Community Reserve." "
In
the Motaba River area
in
321
their Conservation
Map
Congo
16.5 Great
ape distribution
in
10E
12''E
-"T
chimpanzee observed 1983-1995
Central
CENTRAL AFRICAN REPUBLIC
Central chimpanzee
Western lowland
^.Ijl Eastern chimpanzee
Western lowland
gorilla
de la Sgogh
(NouabaW-Ndoki NP)
Trinationale
ape species observed 1996-2003
Western lowland
gorilla
bOMASSA
f^S
"TRtANGQE
'
'
lilLIKOUALA
'^
')
<
GOUALpu|o--M^
,.,.
'^'^G/^.
/^ E Q U A T O R I A^L
gorilla
^/.j^ Central chimpanzee
chimpanzee observed before 1983
Unidentified
le-'E
Estimated range
--*--i..i'
Confirmed range
Species
Central chimpanzee observed 1996-2005

Central
16'E
f"
country profile
this
-^
LIKOUALA
7^ SWAMI
SWAMP
I
t
'
tnpfondo
T, "Guinea
2N
mont-
y"
T^
\S
\\
Lacias
Lac
/ Liksâlaî aux-Herti^
,
Community Reserve
0"N
2"S
*i^Bambama-/
"X
,-
^.
'!.'>.
'
'
-A.
i""
POOLE-'-
DEMOCRATIC
-^
5s6/.m6a/a.
L6kana
NP/
LeflnV,
"^
REPUBLICOF
C-'x__
r^xl
THE CONGO
V r r
Conkouati-'-',
Douli
4'S
NP
HELP
.Dimonika
"^J^'-A,
Chimpanzee
^%>V
Sanctuary
^\
Tchimpounga
Chimpanzee
Sanctuary
Sanctuary/
.;->.
Pom(e-.Wo/re
.^
/',
^^
O
,^-\
Kinkala
Brazzaville
-^
4'S
Madingou
Loubomo
'%
t
'^
Lesio-Louna
Reserve
BR
~^-
KOUILOU BASIN
CABIN DA
ANGOLA
10E
322
300 km
200
'
O
14E
16'E
18E
6S
AFRICA: Congo
northeast Congo, a density of 0.3/km^was found."
The density
southwest Congo was found
in
lower than that
and
NP
the Nouabale-Ndoki
in
north, but about the
same
as
in
be
to
the
in
Equatorial Guinea
year
come from
Both gorilla and chimpanzee populations

are threatened by
human
conflict, the
Congo
in
expansion
of
and associated hunting.^'" The Ebola virus
major threat
and a
in
Gabon
the border region with
gorilla population
was wiped
out at Lossi,
in
community-protected forest some 50 km southwest

" The population in the park
of Odzala-Koukoua NP.
itself
were also believed
to the city
Mayombe
the southwestern
towns and
in
for
Congo and neigh-
cities in
was feared
to
be under threat
in
Figures from the markets

that
for
Brazzaville indicate
in
and chimpanzee carcasses account
gorilla
number
about 2 percent of the total
animal
of
carcasses, and 2.23 percent of the total weight of
meat
and chimpanzees are also
for sale.' Gorillas
frequently killed or
maimed
by traps and snares
intended tor other forest animals such as antelope.'
Some
2004."
local traditions involve great apes.
The
Kwele, Kota, Mboko, and Djem ethnic groups
The
In
effects of civil
the
civil
number
efforts
war and
killed
were hindered as a
meat
raided for
since closed.
to feed
Most
and conservation
result of the overall
The Brazzaville zoo
hungry people, and has
The John Aspinall Foundation IJAFI

Institute (JGIl airlifted the
and the Jane Goodall

gorillas
unknown
aftermath, an
its
apes were
of great
and chimpanzees from the zoo
of the fighting
occurred
in
conflict reaching the
forests
where apes are most
that occurred during the
helped protect
of
In
powdered chimpanzee and
to
gorilla
improve strength and
chimpanzees
in
in
is
likely to
the Motaba region of northeast Congo
against eating their meat;

that
tribal tradition states
Vili
chimpanzees are reincarnations
Continued commercial logging
Disease
in
1992,
in
late
now
in
Congo and Gabon since the
human
eating
in
an estimated 0.01 gorillas and 0.02
chimpanzees per square kilometer per
year, which
represented 5 percent and 7 percent of their pop-
rate,
was higher than
These numbers represented

only;
the re-
and so considered unsustainable.'"

local
consumption
market trade was unquantified. Hunting,
cluding with the use of automatic weapons,

nized as a major
management problem
in
is
in-
recog-
the Lac
Tele/Likouala-aux-Herbes Community Reserve."
In
the 1980s, staff at the Brazzaville gorilla orphanage
estimated that 400-600 gorillas were
killed
each
hunting as a
1990s, and continues to spread. The disease
result of the victims
every part of the area."
rivals
Congo." The Ebola virus has
has claimed over 140
placement
people.
of
Conkouati-Douli
anzee meat taboo.
Aka (BaAka or Bayakal men were
ulations respectively. This
in
NP, and the consequent substantial immigration,
recorded as eating chimpanzee or gorilla meat, and
killed
NP
the Poole region are protected by local taboos
occurred sporadically
Hunting
Hunters
Conversely,
health.^"
have
the country's wildlife during
hunting of apes occurred
bones and hair
and around Conkouati-Douli
Ebola hemorrhagic fever
of
other parts
are traditionally added to the bath water of babies
threat to apes
40 percent
some
with those of cultures that do not consider chimp-
had been
this period.
In
young men.'"
Congo, such as the Odzala and Ndoki regions,
war would have further
accessible. This
much
of
however, has diluted these indigenous beliefs
reduced travel to those forest regions that
more
cision ritual for
of infrastructure
abundant. The general breakdown
previously
1997.
in
the south of the
little
country, however, with
more remote northern
in
north Congo eat gorilla meat as part of a circum-
war
disintegration of law and order
was
to
region.'''
boring countries as urban incomes have increased.
THREATS
is
of the gorilla parts
There has been an increased demand
bushmeat
logging,
may have
northern Congo, although this
and orphans brought
Gabon.'"
in
in
been an underestimate.'^^' Many
primates.
reported
Gabon;
in
late 2002,
In
in
the north of
lives,
probably as a
meat from
an outbreak
infected
Ebola was
of
Congo on the border with
some areas more
than 90 percent
of
the
populations of gorilla and chimpanzee were killed

during this single episode. The epidemic appears to
have
killed all but
seven
of a
study population of 143
gorillas in the Lossi Gorilla Sanctuary, for example.
An Ebola outbreak has not yet been confirmed in the

nearby Odzala-Koukoua NP,"" but by September
2004, an 80 percent decline
in gorilla
sightings
in
the parks Lokoue Bai sparked fears that Ebola
had arrived
"
Other diseases reported
in
apes
in
Congo
323
World Atlas
under which two Biosphere Reserves have been
NP and Dimonika
designated: Odzala-Koukoua
the
in
Mayombe mountains.
The main laws dealing with
wildlife conservation
and use are Law 48/83, which protects gorillas and
chimpanzees,'"" Law 49/83

Decree 85/879
November
1983, and
of April 21
1985.
Law 16/2000
of
20 2G0G establishes the Forestry Code,
A newly released
which aims
chimpanzee
forest
at
July
of
achieve sustainable
to
ecosystems
management
Congo. The Ministry
in
Conkouati Lagoon
Economy and
takes a last look
sible for wildlife conservation
back.
including
the Environment IMFEE)
management
of
of
of Forest
is
respon-
and regulating use,
protected
The
areas.
Congolese government has recently banned the

include yaws, a debilitating disease closely related
to syphilis, with 21*
from a sample
"
being infected.
production of shotgun cartridges.^
Congo
420 gorillas
of
is
part
Congo Basin Forest
the
of
Partnership (CBFP), an international
initiative inclu-
ding governments, nongovernmental organizations
(NGOs), and the private sector, which
Habitat loss
It
is
estimated that 170 km' of forest was cleared
annually
about
Congo from 1990
in
percent
0.1
to 2000.
representing
per year." Timber
of the total
is
to
is
"committed
conserving the forests of the Congo Basin while
promoting sustainable economic and social de-
velopment
in
the region."
also selectively exploited wherever dryland forest

is
accessible. Selective logging can
habitat for gorillas,
supplies
which can often
secondary
in
forest,
but
improve the
find
good food
associated hunting reduces gorilla numbers.'''

of the forests of the north
so far because
is
of
to logging
poor road infrastructure, but
management
companies."
of the
units have
In
bushmeat hunting by
relatively
to
forest wildlife,
bushmeat
local
to
this
remaining pris-
been assigned
Congo, roads established
and maintained by logging concessions
access
Some
have remained unlogged
changing and today many
tine forest
the
reality,
in
intensify
Protected areas
There are three main categories

Congo: national parks (four
(six sites),
and hunting reserves
(four sites)."
in
There
number
are also four faunal sanctuaries and a
of
other reserves.'* The existing protected area net-
work covers an estimated
percent of the country's
land area. The Congolese government has recently
launched an evaluation
areas, with an
of the country's
protected
emphasis on assessing gaps between
parks, with support from the Wildlife Conservation
populations of
Society (WCS), and aims to create a national service
unexploited
lowering the cost of transporting

in
market among logging
many cases, offering

company workers."
for protected area
anzees are found
and
Western lowland
management."
wildlife
gorillas
in all
NP
Nouabale-Ndoki
and central chimp-
three national parks. The
covers an area of 4
contains about 2 percent of
all
93 kml^
It
Congo's forests. Most
International legislation
of the
Congo has
or acceded to the Convention on
logged once, at a tree intensity of about 100/km',
Biological Diversity (19931, the Convention on Inter-
and active logging concessions largely surround the
ratified
national Trade
in
Endangered Species
Fauna and Flora ICITESl

to
Combat
(1983), the
Desertification
UN
(19991,
of
of
Wild
Convention
the
African
periphery
'"
park.^'
Bomassa
to
Human
In
the west and south has been
impact on the park and
buffer zone
administers the park
Nature and
2001,
the
is
in
collaboration with the
Odzala-Koukoua
expanded
Reserve and M'boko Hunting Reserve (both

south
to
the
WCS
MFEE.
NP was
Natural Resources (1981), and the World Heritage

in
in
considered minimal."
Convention (1987). The country also participates
UNESCO's Man and Biosphere (MAB) Programme
324
area
faunal reserves
providing hunters with greater
market and,
LEGISLATION
of protected
sites),
include the Lekoli-Pandaka Faunal
of the park)
and an enormous area
to the
of forest
AFRICA: Congo
east, previously assigned for
west and
to the north,
timber exploitation. The park now covers 13 456

knnôn the northwestern edge
watershed, with altitudes from 300
human
the absence of
tends
more herbaceous
protected
areas
Congo, Cameroon, and the
in
(see
CAR
country
many
clearings
attractive
(swampy
of
rain
bais] that
mammals,
large
to
lowland
western
of
(mean
gorillas
and chimpanzees (mean 2.2/km'|
5.4 /km^l
In
Central Africa.' Wherever gorillas and elephants
lations of
gorilla habitats,
of the
Similarly, there
is
Congo, Angola, and DRC."
NP
has been
the
European Union program. Conservation and RatioForest Ecosystems
of
in
Central Africa
(ECOFACI, which has brought about
declared a Biosphere Reserve
Conkouati-Douli
coast
5
kml"
common
is
in
NP was
in
Congo and covers
terrestrial part of the park Includes several active
logging
villages
'ecodevelopment zone'. Hunting has proved

to
significantly to
Noire,
and
the
bushmeat market
of
has severely affected large
this
It
an
difficult
Polnte-
mammal
was managed by lUCN-The
populations. The park
World Conservation Union

since which time
In
area contributes
control, since the Conkouati
for five years until 1999,
has been managed by WCS,
in
in
the LIkouala
swamp
both gorillas and chimpanzees.

Involved
In
region supports
WCS
is
becoming
management of this reserve, desRamsar Site In 1998 and owned by
the
ignated as a
the local communities. Finally, the LossI Gorilla

a small reserve effectively created by
Sanctuary
is
the
community, who established an eco-
local
tourism project here.
It
Is
taken up by national
inclu-
seems
part of the forest
to
have been particularly
affected by logging.
Conservation
field projects
developed
at several sites
Odzala-Koukoua
WCS
being
is
around Congo, including
NPand Nouabale-Ndoki NP
has been involved
servation projects
in
with the
MFEE and
des Bois
(CIB),
periphery
of
in a
number
Congo, including
of
con-
project
the Congolaise Industrielle
a logging
company, around the
Nouabale-Ndokl NP. A set
of
agreed
guidelines for hunting has been developed, which
includes a ban on the hunting of apes and other

endangered species and the export of meat from
the concession. As a result, commercial bush-
meat hunting has been reduced, and

ported
in
1999 that the incidence
it
was
of gorilla
re-
and
chimpanzee hunting had dropped by an estimated
MFEE.
The Lac Tele/Llkouala-aux-Herbes Community

Reserve
If
would be an extensive area
ding fully protected and limited-use zones. Congo's
to
the northern part of the park. The
concessions and many
this
Ecotourlsm, Including gorilla watching,
1977.
situated on the Atlantic

of
governments,
significant
Gorillas and chimpanzees are said
045
be
NP
southwest
the
in
poaching. The Odzala-Koukoua
In
Mayumba
the
area, to include parts of
activities
of
to
an emerging proposal for
Mayombe transboundary
1997 through 1999 when conservation
administered since 1992 with the support
NP
Gabon.
in
were reduced, the Odzala-Koukoua
NP
connect Conkouatl-Doull
NP
war from
civil
of a trans-
neighboring Gabon. The government also plans
'"
Apart from the period during the
Congo
of
Bambama-Lekana
boundary protected area with Bateke Plateau
to
herbaceous vegetation.
of
January 2004, the government
NP, which contains chimpanzees, as part
through seed dispersal as well as
to
up the forest and encouraging the
by opening
In
announced the creation
In
would be expected
western lowland gorillas and central
chimpanzees.
Improve
coexist, the latter
decrease
in
The core
profile).
protection zone comprises the national parks of
including gorillas. Odzala has the highest recorded
densities
Use
managed
Central African Republic (CARl are
common
which
In
Nouabale-Ndokl (Congo), Lac Lobeke (Cameroon),
particularly
nal
Sangha
a partner in Trinatlonale de la
and Dzanga-Ndokl (CAR), which together cover

about 7 300 kml All support important popu-
forest contains
growth
is
(TNSI, a transborder conservation zone
open-canopy Marantaceae forest with
of the forest of
dense undergrowth. The northern area
are
Congo
Odzala
Over 90 percent
71.
fairly
is
is
such as t>/larantaceae species (see
vegetation,
Chapter
to
influence, gorilla density
be higher where there
to
Congo River
600 m.^ " In
of the
Transfrontier initiatives
also used for research.
90 percent over a two year period."
in
addition, the
company gave up a large part of its concession

an area where there are apes with little fear
in
of
humans, presumably because they have not had

previous contact with people. The World Bank
has become Interested
as a
model
for
in
Africa.'^' In addition, the
now
requires
all
promoting the project
other concessions
In
Central
Congolese government
logging companies operating
In
325
World Atlas
pay for ecoguards' and wildlife
northern Congo
to
management on
their concessions.'
Research
the
into
Congo has taken place

an
1
ecology, and
chimpanzees
at a
number
in
.3
swampy
open,
forest
clearing
covers
that
km", and the Goualougo Triangle, both located
Nouabale-Ndoki NP;"Maya Nord
Sanctuary,
Gorilla
and
area;'''
km
50
in
Odzala-
faa/in the
Koukoua NP;" the Likouala swamp

Lossi
of great
reintroduction projects
Aspinall Foundation
southwest
Lefini
in
IJAFl
In
or-
gorilla
was established adjacent
1994,
In
Faunal Reserve by the JAF
in
to the existing
partnership with
The area was
be
to
southern
for gorilla rehabilitation. In 2004, the
was combined
with the
Lesio-Louna Reserve, creating a protected area
kml There
700
are currently 23 gorillas
project's care, 15 of
in
of
the
which have been successfully
reintroduced. There has been one birth
the re-
in
introduced group.'
opened
km^
in
km
50
it
is
is
north of Pointe-Noire,
at least
numbers
of
was
run by the Jane Goodall
the largest sanctuary of
houses
It
increasing
lies
1992, and
Institute IJGII:
in Africa.
in
primatologists and
held
March 2003 under the GRASP
Brazzaville
in
initiative to
discuss
Congo. A
in
ape survival plan (NGASPl was
produced." The
mam
working groups
at
recommendations from the
summarized
the meeting are
Increase effectiveness of surveillance and

antipoaching brigades
kind
its
115 chimpanzees, with
orphans
most
arriving. In
forest concessions,
in
create a national antipoaching brigade, establish
an interministerial committee
combat poaching, and
among
conservation of wildlife
police,
help
to
publicize the law on
the army,
customs, and courts.
Ensure
effective
surveillance
protected
of
areas, including better supply of equipment,
some
designate
areas as World
protected
Heritage Sites, and promote transborder
agement
man-
of great apes.
Develop
system
monitoring
for
illegal
concerning great apes and en-
courage the national networking
of environ-
mental information management, with an
emphasis on great apes.

Undertake research
bution,
and biology
including
their
the status, distri-
into
of great
apes
in
Congo,
dynamics and
population
cases, the arrivals are young chimpanzees confis-
diseases, develop national research into great
cated by the Congolese authorities from hunters
apes, and establish a database of
trying to sell
them. The center also engages
local
in
all scientific
studies of great apes.
community development work and environmental
Develop a national policy on tourism, promote
awareness programs.
tourism
A Congolese NGO
called Habitat Ecologique et
Liberie des Primates (HELP), established
in
1991,
also cares for young chimpanzees confiscated by
the Congolese authorities. Since
November
1996,
36 orphan chimpanzees have been released

the
forests
of
Conkouati-Douli
NP
into
(previously
Conkouati-Douli Faunal Reserve). To monitor the
success
field
of the introduction, a
team
of
Congolese
workers tracks the chimpanzees each
day. Of
20 animals released, 14-17 survived and readapted

to life in the wild.'"
HELP
no longer rescues
orphan chimpanzees, and the release
326
was
activities
The Tchimpounga Chimpanzee Sanctuary

covers 73
of
other conservation experts
draft national great
John
1986, the
started a
the grounds of Brazzaville zoo.
sector of the Lefini Reserve

An international meeting
ape sanctuaries and
Congo.
the Congolese government.'"
used
process (see
reintroduction
of
Lesio-Louna Sanctuary (now gazetted as a
reserve!
the
of
4.51.
below.
There are a number
the
Box
conservation action tor great apes
Sanctuaries
in
assessment
at
Odzala-Koukoua NP.^
phanage
Post-
for 2003.
mostly
of sites,
the north of the country. These include Mbeli Bai,
in
was planned
release monitoring will continue until 2013 to allow
population,
social behavior of gorillas and/or
suitable candidates
of
new
the last
create
among departments
incentives to
of
promote
government,
local
tourism
throughout the country, and carry out a feasibility
study of ecotourism
in
areas inhabited by
great apes.
income
for
communities such as agroforestry,
fish
Develop alternative sources

rural
of
farming, and beekeeping.

In all forest
concessions
in
Congo, promote
the results of the joint project of the
and the
WCS,
CIB,
IvIFEE.
Create an autonomous wildlife and protected

area
management agency
AFRICA: Congo
Undertake both education and conservation-
and wild apes." A second workshop was held
awareness
Brazzaville
activities.
healthcare, through the support of the

Ministry of Health and Population, and
Particular concerns over the threat of the Ebola

virus to great ape populations
been raised and
calls
in
Congo have also
made
have been
in
March 2003 on Ebola and preventative
in
workshop shared understanding
of
MFEE, the
ECOFAC. The
the virus and
started to fornnulate a strategic approach to deve-
for increased
research on vaccines and ways to vaccinate people
loping preventative
measures and further research.
FURTHER READING
Bermejo, M. 11999) Status and conservation
primates
of
in
Odzala National Park, Republic
of the
Congo. Oryx 33
I'll:
323-331.
Blake,
Rogers,
S.,
Ecotogy33
Bowen-Jones,
E.,
Fay, J.M.,
E.,
Ngangoue,
Pendry,
S. 11999)
The threat
this threat could be diminished.
Fay, J.M.,
Ebeke,
M.,
Swamp
G. (19951
gorillas in northern Congo. African Journal of
285-290.
131:
Oryx 33
Agnagna, M. (19921 Census
apes and other animals from the bushmeat trade
to
(3):
in Africa,
and how
233-246.
northern Republic of Congo. American Journal of Prlmatology 27:
of gorillas in
275-28/..
Kano,
Asato, R. (1994) Hunting pressure on chimpanzees and gorillas
T.,
Congo. African Study Monograpfis 15

Leroy, E.M., Rouquet,
Swanepoel,
P.,
Formenty,
P.,
Souquiere,
R., Zaki, S.R., Rollin, P.E.
in
the Mobata River Area, northeastern
U3-162.
131:
S.,
Kilbourne,
A.,
Froment, J.M., Bermejo, M.. Smit.
S..
Karesh, W.,
(2004) Multiple Ebola virus transmission events and rapid decline of
central African wildlife. Science 303: 387-390.
MFEE
pour
(2003) Rapport final de I'atelier sur ielaboration d'un plan national
la
survie des grandes s/gnes.
GRASP:
Great Apes Survival Project. Ministere de I'Economie Forestiere et de lEnvironnement, Republic of Congo.
Nishihara,
(1995) Feeding ecology of western lowland gorillas
T.
Pnmafes 36
Walsh,
(21:
Abernethy, K.A., Bermejo, M., et
P.,
in
the Nouabale-Ndoki National Park, Congo.
151-168.
at.
(2003) Catastrophic ape decline
in
western equatorial
Africa.
Nature
422:611-614.
Wilkie, D..
Shaw,
E.,
Rotberg, F, Morelli,
Basin. Conservation Biology
(61:
Auzel,
G.,
P.
(2000) Roads, development, and conservation
in
the
Congo
1614-1622.
MAP DATA SOURCES

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

Stevens,
E.F.,
Arluke.
A.,
In:
Beck, B.B., Stoinski.lS., Hutchins, M., Maple,
eds. Great Apes
and Humans: The
Etfiics of
T.L.,
Norton,
B.,
Rowan,
A.,
Coexistence. Smithsonian Institution Press,

With additional data by personal communication from Maisels,
Goossens,
Poulsen,
Setchell,
B.,
J. P.,
swamp
12004), Poulsen, J. (20051, Ron,
J.
(2003)
Home
in
free.
BBC
Wildlife 21
(2003), Stokes, E.
30-35.
[21:
Clark, C.J. (2004) Densities, distribution, and seasonal
forest
in
movements
northern Congo. International Journal of Primatology25
Various authors (2003) Annotated
team
F.
and from the following sources:
(2003),
Brazzaville,
map
Congo on
prepared by
UNEP-WCMC
and modified
12):
of gorilla
and chimpanzees
in
285-306.
at the
meeting of the national
GRASP
Emma
Stokes
April 2 2003.
ACKNOWLEDGMENTS
Many thanks
(Wildlife
to
Amos Courage
[John Aspinall Foundation), Ian
Conservation Societyl for their valuable
comments on
Redmond
[Ape Alliance/GRASPl, and
AUTHOR
Nigel Varty.
UNEP World
327
World Atlas
REPUBLIC OF COTE D'lVOIRE

Ilka Herbinger, Christophe Boesch,
Adama Tondossama,
AND Edmund McManus

The Republic
bordered by Liberia and Guinea

Burkina Faso
to the north,
Is
per year, and mainly
and
and the
to the east,
of
was estimated
be 17.3 million, growing at about
to
Africa,
has a land area
It
2004, the population
In
West
in
to the west, Mali
Ghana
Gulf of Guinea to the south.
318 000 km'.
Cote dlvoire
of
2.1
percent
the southern coastal
living In
region and the main commercial center of Abidjan.'
Cote d'lvoire gained independence
1960,
In
from France, with
Houphouet-Boigny as presi-
Felix
dent and chairperson of the single ruling party. This
arrangement lasted
until 1990,
system was established.
elected and remained
The economy
a multiparty
first
contested
Houphouet-Boigny was
presidential election,
1993.^
when
the
In
In
office until his
re-
death
in
varus].
It
great ape
of
found
Is
Cote
in
western chimpanzee [Pan troglodytes
has historically been widespread through-
out the country, with
most
individuals living in the
16.61.
Surveys
(Map
rain-forest zone
and 1989-1990" suggested a
11000-12 000
chimpanzees
with
Individuals,
living
1986-1988"
In
population of
total
over
half
the
protected areas, including
in
national parks (NPsI and reserves, and the remain-
der
living
poorly protected
in
or unprotected areas.
In
classified
forests
2003, the population size
was estimated
to
which according
to the figures in the
other country
some 21-36
percent of the
be 8 000-12 000 Individuals,"
would add up
profiles,
total population of
to
western chimpanzees.
dominated
Sites that are thought or confirmed to hold
by agriculture and related services, which together
Important numbers of chimpanzees include the TaV
engage about two

one
historically
exporters
Cote d'lvoire
is
thirds of the population.
of the world's largest
of coffee,
economy
its
of
It
is
producers and
cocoa beans, and palm
oil,
so
sensitive to international prices for
is
these foodstuffs. These prices were high
in
the mid-
1990s, which, together with other factors such as
the
petroleum resources,
discovery of offshore
allowed international debt
Annual growth
about
in
percent
to
be accumulated.
gross domestic product (GDPI
in
was
the period 1996-1999.
This period of relative prosperity

to
was brought
leading to Instability, counter-coups, and a
brokered
with
Community
of
France began
into 2003,
the
civil
of
the
Economic
West African States (ECOWASI and

to
further troubles
take effect. Although there were

in
Comoe
2003, the agreed peace process
continued and several thousand French and West
Bandama, Bossematie, and

Forest." "
The high proportion
most
fact that
in
West
in
2004 by a United Nations peacekeeping
However, an upsurge
in
violence
In
November
2004 slowed down the ongoing peace process, and

triggered a
UN embargo
either side.
The economy contracted each year
on arms dealing with
once the
Africa - have been heavily logged,
and deforestation
is
now widespread
outside the
protected area system.

It
has been estimated that there are around
4 500 chimpanzees
and up
to
In
the TaT
500 Individuals
In
NP
10.4-1. 7/km'l,
the nearby N'Zo
Faunal Reserve and Haute Dodo and Cavally-Goln

Classified Forests."
for
"" The
last
published census
Mont Peko NP (1989-19901 estimates
a total
population of 78," but a later census suggested a

significant population of
replaced
chimpanzee
of the country's forests -
There might
force.
of the
population living within protected areas reflects the
the disarmament, demobilization, and rehabilitation
combatants. The West African contingents were
the nearby Songan-
Tamin-Mabl-Yaya complex and Monogaga Classified
African troops remained to keep order and facilitate
of
Nature Reserve,
Strict
NP, Marahoue NP, and the classified forests
Haute Dodo, Cavally-Goln, Haut Sassandra, Haut
of
war,
when peace agreements
support
NP, N'Zo Faunal Reserve, Mont Peko NP, Mont
Sangbe NP, Mount NImba
largest
an end by a military coup on December 25 1999,
which continued
328
Only one species

d'lvoire, the
the
Is
still
classified
about 320 chimpanzees.'
be about 400 chimpanzees
forest
of
connected by forest corridors
NR" Mont Sangbe NP
Mont Peko
to
might hold
recent survey.'"
Mount NImba
between 2000 and 2004, and the outlook was poor
Is
for 2005.
d'lvoire section."
thought
to
Strict
least
at
chimpanzees,"' or up to 260 as estimated
which straddles Cote
In
Haut Sassandra, which
in a
55
more
Nature Reserve,
d'lvoire, Liberia,
and Guinea,
have about 59 chimpanzees
in
the Cote
The whole reserve, together with
AFRICA: Cote
Map
16.6
Chimpanzee
>ri
distribution
in
Data sources are provided
Cote d'Ivoire
end of
this
country profile
4'W
6"W
8"W
at the
d'Ivoire
^--,
Species
r~'
'
Western chimpanzee observed 1983*1995
Cburkina
FA S
''
Western chimpanzee observed 1996-2003
MALI
^j
1983
Western chimpanzee obsen/ed
'J
Western chimpanzee observed before 1983
Western chimpanzee presence alleged
Western chimpanzee
after
locally extjnct since ca.
19
Confirmed range
Western chimpanzee
[
Estimated range
10'
I^7/j^
V-
GUINEA
.....
.
^Mont
..
'^^TS
'
9^^-i
r "^^^
C6TEîr>VOIR
C6TEUjrv>VOIR
';
(>^
(;>
'^
O/^
O
\ ( L,
/^ $ S \
1/
Bandama'
<^
Mankono
Haul
J--
^r
Sangb6NP\'"-v
Sanqb6NP, "-t_
Oj 6l\
Western chimpanzee
9''
i-^M.-i-"*^
,'
\>--i''^
Bondoukou
'
8"N
^VaTOua^^
i_JcJ
J
\
i
$
^Zttenoula
'v'
.,
Sakassot,
a^j^'**
Makû^Â Wo
*\.BouafIe
^
Li
'^
^^
^-.Q
FaR
WHS J
>
^^
(^
I
/^
(O*
\,
'n'Zo Partial
\J
\^ Mbahiakm
V'
'^'Agbovllle
,'
<^
GHANA
AgpjbHekrou
DaQlrJ^
dÂ a-Ô^J^'
O^/(gt0W/te
a
I
GagAra-^%^fc VJ '-'ft^^
GagW/â.*.
U-'-'l^
^V /s
Ci)
z^''-
Vaya Complex CF
^^
ntr
Lug"'
I
Gulf of Guinea
TLANTIC OCEAN
4"N
300 km
329
World Atlas
Chimpanzees are hunted

for
Chimpanzee meat
crops.
and
meat, for ingredients
for
medicine, and because they raid
traditional
sold
is
urban markets
in
However, taboos against
village restaurants.''
consuming chimpanzees
killing or
out Cote d'lvoire.
through-
exist
among
Religious taboos exist
Muslims, as Islam forbids the consumption
meat
of the
and cultural taboos
of primates. Traditional
mostly stem from legends connecting chimpanzees
and humans: these
formed
ing
humans
and times
in
being trans-
of
assist-
during sicknesses, accidents, births,
war Local
used
traditions could be
promote the conservation
A chimpanzee
humans
of
tell
chimpanzees and chimpanzees
into
to
chimpanzees.
of
Zoonotic diseases including monkeypox and
TaV
National Park, Cote
Ebola are a threat
d'lvoire.
diseases introduced to the wild population through

"r
Herbinger/Wdd Chimpanzee Foundalion
adjacent classified forests, migfit contain over 300
NP was
chimpanzees. Comoe
estimated
NP had
about ^70 individuals." Marahoue

highest density of chimpanzees
estimated
to
be 6.Vkm',
Cote
in
the
d'lvoire,
population of
w/ith a
individuals 11989-19901."
fiave
to
rapid census
A07
1998,
in
however, found only one nest and had only four
chimpanzees, confirming
auditory contacts with
died.'
for
Haut Bandama was estimated
the
may
nearby Songan-Tamin-Mabi-Yaya
complex
hold a combined total of 600 individuals."
may
Approximately 100 chimpanzees

the
to
Bossematie and
of
Monogaga
still
of
chimpanzees
in
Cote d'lvoire
management
of
these mostly isolated populations.
depend upon the
will
THREATS
The southern
half
Cote
of
supported about 160 000 km^
d'lvoire
of
rain
originally
but
forest,
immigration and agricultural expansion resulted

in
widespread deforestation. By 1966, annual de-
forestation
figure that
was estimated at about

was maintained into the
declined to about 2 650
from 1990
to
km'
(3.1
2000. Only 71 170
mained by the end
000 km^'^ a
1980s.
later
It
rapidly,
percent.' tôre recently, the bacterial disease an-
thrax
killed
Other threats
chimpanzees
in
to
in
eight
least
first
chimpanzees
TaT
in
time that this illness was
have affected great apes in a rain-forest

" This infection was tentatively attributed
to
deforestation
the
has
that
allowed
cattle
transport routes from Mali and Burkina Faso to
pass close
to the park,
so that the chimpanzees
may
have caught the disease from passing livestock.
LEGISLATION
Cote d'lvoire
is
much
of
it
in
the
a signatory to the Convention on
Biological Diversity, the Convention on International
Endangered Species
Trade
in
Flora
(CITES),
the
Desertification.
In
UN
Cote d'lvoire
Wild Fauna and
Convention
1995, the
government adopted
national environmental action plan

is
Timber
Combat
to
national strategy for parks and reserves;
in
1996, a
was created and
currently being implemented.
Present
chimpanzee-related
include the
Autonome pour
National de
Ta'i
which
in
offers,
among
populations
in
the
in
Cote
other
activities,
chimpanzee com-
the southern part of the
mental education projects
anzee
of
Conservation du Pare
la
a guided visit to a habituated
munity
conservation
Ecotourism Project
Chimpanzee Foundation (WCF)
bushmeat, and disease.
of
Agreement, and the
Projet
of forest re-
from an estimated 100 000
to
at
NP, which was the
km'
early 1960s to 8 000-12 000 by 2003."
include hunting, trade
1999, an epidemic of acute respiratory
In
projects
degraded. Chimpanzee numbers have declined

even more
Ebola
community under study
of the
disease reduced that community by a further 25
percent) per year
of that period,' with
suffered
of the
exist in
" The future
Classified Forest."
NP
Tai
in
new subtype
of a
and 25 percent
setting.'
be 300." The classified forests
330
virus,
known
chimpanzees
chimpanzees
from an infection
had greatly declined.'^ The
population of
these small populations, as are
interactions with farmers, hunters, researchers, or

tourists. In 1994,
their presence, but suggesting that the population

total
to
Ta'i
NP. The Wild
carries out environ-
the vicinity of chimpd'lvoire,
including
AFRICA: Cote dIvoire
interactive theater plays,
presentations.
WCF
is
a national biomonitoring
ting
Tai
to
to
WWF-The
it.
main
The
country's
In
chimpanzees
TaT NP,
on
project
under the supervision
Boesch, and focuses on various aspects
in
and enlarge the two forest
Conduct a survey
chimpanzee population,
the
meat hunting
Gulnean
Christophe
of
the nearby populations
to
enforcement and surveillance
also being carried out
is
Cavally-Goin and
NP
to
Haul
Sassandra Classified Forest. Improve law
Surveys.
research
field
of
these Cote d'lvoire
corridors that link Mont Peko
Global
WCF activities.
to various
link
to
Liberia. Protect
Conservation Organization has provided financial
support
NP, and
chimpanzees
estimate the
Cote dIvoIre and to iden-
In
immediate threats
tify
Implemen-
in
program, including
methods
the development of better
newsletter, and film
also involved
In
belt,
efforts.
the current
of
and bush-
habitat,
the largest forest block
which includes
Tai
Faunal Reserve, Cavally-Goin, and
of their
behavior, ecology, cognition, and genetics, and on
the
In
NP, N'Zo
Haute
Dodo. Repeat a nationwide survey, including
disease transmission between chimpanzees and
Marahoue NP, Comoe NP, Mount Nimba
humans.
Reserve,
and
Sassandra
Haut
the
and
Songan-Tamin-Mabi-Yaya complex. This

Some of the priorities for conservation
geting the
chimpanzees
of
allow the rate of population decline
in
rural and
urban areas have
been suggested. Including

Protection. Assign a higher protected area
NZo
and
Forests.
Create as
Faunal Reserve and
particular,
create
corridors
link
to
In
program
of
the center of Abidjan
should play a crucial rote
in
such education
programs. Ecotourism development could
protected forest
chimpanzee populations.
Banco NP
areas.
also be initiated
corridors as possible to link the country's
fragmented
regular visits of school classes to protected
Haute Dodo Classified
many
be
Education. Education and awareness-raising
Cote d'lvoire are outlined
campaigns
Cavally-Goln
will
estimated.
action tar-
here.
status to the
to
Tai NP,
In
the
in
promising sites such as
Mont Sangbe NP, or the Monogaga
Classified Forest.
FURTHER READING
Chatelain, C, Gautier,
L.,
Spichiger, R.A. 119961
Biodiversity and Conservation 51
Formenty,
P.,
recent history of forest fragmentation
Boesch, C, Wyers, M., Steiner, C, Donati, R, Dind,
outbreak
among
wild
in
southwestern Ivory Coast.
37-53.
chimpanzees
living in a
ram
F.,
Walker, R, Le Guenno, B. 11999) Ebola virus
forest of Cote d'lvoire. Journal of Infectious Diseases 179
ISuppl 11:5120-5126.
Herbinger,
In
I.,
Boesch, C, Rothe, H. 12001) Territory characteristics
among
three neighboring chimpanzee communities
the Tai National Park, Cote d'lvoire. InternationalJournal of Primatology 22
Kormos,
R.,
Boesch,
C.,
Bakarr,
ll.l.,
12):
l/i3-167.
Butynski, T.M., eds (20031 West African Cfiimpanzees: Status Survey
and
MarchesI,
P.,
MarchesI, N., Fruth,
Pnmates
B.,
Boesch,
C. 11995)
Census and
distribution of
chimpanzees
in
Cote
d'lvoire.
36: 591-607.
MAP DATA SOURCES

Map
16.6

In:
Kormos,
R.,
Pan
Boesch, C, Bakarr,
following sources:
troglodytes: taxonomy, distribution, abundance, and conservation status.

M.I.,
Butynski, T.M., eds. West African Ctiimpanzees: Status Survey
and
Conservation Action Plan. lUCN/SSC Primate Specialist Group. lUCN, Gland, Switzerland, pp. 5-12.
Herbinger,
eds.
1.,
Boesch, C, Tondossama, A. 12003) Cote
d'lvoire. In:
Kormos,
R.,
Boesch, C, Bakarr,
West African Cfiimpanzees: Status Survey and Conservation Action
Plan.
M.I.,
Butynski, TM.,
lUCN/SSC Primate
Specialist
Group. lUCN, Gland, Switzerland, pp. 99-109.

With additional data from:
331
World Atlas
Fischer,
F.,
Gross, M. (19991 Chimpanzees
Hoppe-Dominik,
Comoe
the
in
and status
B. 119911 Distribution
Pan
National Park, Cote d'lvoire.
Newsi
Africa
chimpanzees [Pan troglodytes varus] on the
of
121:
Ivory Coast.
19-20,
Primate
Report 31:^5-57.
Marches!,
P.,
Marches!,
Boesch,
N., Fruth, B.,
C. 119951
Census and
chimpanzees
distribution of
Cote
in
d'lvoire.
Primates 36: 591-607.

ACKNOWLEDGMENTS
This country study draws extensively on the Cote d'lvoire chapter
African Cfiimpanzees: Status Survey
Planck Institute for Evolutionary Anthropology) and
to a further
in
the
Plan."
lUCN/SSC Primate
Thanks
reviewer
Specialist Group's
who wishes
to
West
Hohmann (Max
reviewer Gottfried
to
remain anonymous.
AUTHORS
Ilka
Harbinger, Wild Chimpanzee Foundation
Chrlstophe Boesch, Wild Chimpanzee Foundation
Adama Tondossama, Tat National Park

Edmund McManus, UNEP World Conservation
Monitoring Centre
DEMOCRATIC REPUBLIC OF THE CONGO

Nigel VARPf

The Democratic Republic
merly
Zai're)
of the
covers 2 3^5 410
km^ making
countries
In
independence from Belgium
in
the three
largest
Africa.
General Mobutu as president
was
one
It
of
gamed
It
an army coup d'etat
political
installed
rule
for-
1960, and after a
time of
instability,
in
1965. His
largely unchallenged until the late 1980s,
sinated
sworn
in
In
2001 and his son, Joseph Kabila,
improved democracy,
liberalization. Active
between the government and the main armed
and episodes
Rwanda
This
99^ further destabilized the situation, as President
challenge
Mobutu gave sanctuary and support
Hutu refu-
to
gees from the former Rwandan Army. The

inhabitants living
allied
in
Tutsi
the east of the country then
themselves with other discontented armed
groups and launched a successful attack on the

Zairean
army and
the
Mobutu regime
leader, Laurent Kabila,
was
in
1996. Their
installed as president,
but within two years the political situation
DRC began
to
in
unravel and Congolese
eastern
rebels,
supported by Rwanda and Uganda, seized major
towns
there,
prompting
Namibia, and Chad
to
Zimbabwe,
intervene.
Angola,
By May 1999,
diverse rebel armies controlled large areas
In
the
to the
of
combat
recent
of
in
history
2002
in
coup attempts
place, despite repeated
is still in
in
in
and economic
rights,
groups, creating a transitional government
regime was growing, and there were army muti1991 and 1993. '"'The genocide
human
diplomacy by South Africa and
the United Nations has since achieved a settlement
that
in
was
as president, promising a program of
by which time opposition to the corruption of his
nies
the east of the country.

is
fundamental
conserving the great apes
economic circumstances
in
the
to
DRC, and
of the country's 55
million people, a population that
is
2.9 percent annually according to
growing
at
about
2003 estimates."
The Belgian colonial government created
little
infrastructure or social capital, although at inde-
pendence there was
transport
system that
included strategically interconnecting roads, rivers,
and
railways.' Little
regime after an
more was done by the Mobutu
initial
flurry of
public
1966-1974 (mostly funded by foreign
much
that
had been created was
neglect, underinvestment,
works
loans),
in
and
later destroyed by
and war The country's
abundant natural resources include
cobalt, copper,
north and east of the country, and had begun to
gold,
finance themselves (and their foreign allies]
manganese, tungsten, tantalum, cadmium, petro-
looting
332
and coltan." " President Kabila was assas-
gold,
Congo IDRC,
by
mineral resources such as diamonds.
leum,
diamonds,
timber,
zinc,
and
uranium,
untapped
tin,
silver,
coal,
hydroelectricity
AFRICA: Democratic Republic of the Congo
resources,
yet
domestic
gross
2002-2003,
in
was estimated
product IGDPl
US$5.7
at
was
gross national income IGNI) per person
US$90, or US$0.23 per person per
ttian
ttie
Human Development
was
168tti out of
halt
DRC
177 countries ranked globally'"
percent.'^
Africa's
of
less
" and
day,'
Index position of
Agriculture accounted for 55 percent

industry 11
billion,
of
GDP, and
DRC possesses more
forests." Forests cover
than
broad-leaved
closed
tropical
roughly 15 000 km'
Africa
in
for
Having reviewed
cover.'''
DRC from
1960
to 2000,
Monetary Fund con-
analysts at the International
the south.
in
occupied
It
the early 1990s,'^down from
1959-1960." Most
in
gorillas lived in the
Kahuzi-Biega NP, whose 6 000
km'
spread over
is
separate mountain and low-
land sector, connected by a forested corridor. The

park's population suffered a severe decline
Some
conflict of the late 1990s.
The mountain
to
the
in
sources estimated
few thousand gorillas remained by
only a
percentage
in
in
in
an estimated 21 000 km'
2001.""
only 3.5 percent of the land

of
Lake Tanganyika
that
of the
terms. Permanent crops and arable account
economy
Lake Edward
of
making DRC one
two most forested countries
the political
of
corner
352 070 km' (about 59.6
percent) of the country,''
the north to the northwest
corner
gorilla population
restricted
is
the southern section of the Virunga NP.
It
forms
part of the larger 'Virunga' population, inseparable
from the gorillas
NP
in
the contiguous Mgahinga Gorilla
Uganda and the Volcanoes NP
in
Rwanda. An
in
cluded that poor economic policies, bad gover-
estimated 183 mountain gorillas were resident
nance, and war had contributed to the country's
DRC
economic decline during
this
AO year period, but
that the right policies are
now
being put
pave the way for a restoration
in
place to
economic growth.'
of
in
Mount Tshiaberimu
NP
Virunga
lowland
beringei graueri]
[Gorilla
gorilla
endemic
DRC. The mountain
to
and
beringei]
chimpanzee
eastern
the
troglodytes schweinfurthii] are also present.

not
known whether
gorilla gorilla]
troglodytes]
persist
still
of
The bonobo has
the
Congo
Map
leads
of the
500 000 km'
The
great
Numbers
numbers.'"
about
uncertainty
of
more than
known from many
research efforts within the Salonga National Park

INPI and the
reserve),
Wildlife
Lomako
forest
proposed forest
(a
between the Lomami and Lualaba
along the
Eastern chimpanzees occur
Kasai-Sankuru River,
Research Project
field
in
site,
to
Rivers,
the
Lukuru
at
Wamba,
Sudan border
of the Albertine Rift,
in
the
DRC.
forested areas
in
the
in
Forest
Ituri
in
the eastern regions
Rift,
stretching from the
the north to the southern end of
Itombwe Massif, DRC,
was estimated
total
in
the north-
in
In
the Kahuzi-
total
a study area of
of
100
600 km'." Their
population has been estimated at 70 000-
100 000 individuals.^' This

revision
of
the
is
a significant upward
1980 estimate
of
around 6 200
chimpanzees." However, current estimates are

provisional, as
little
chimpanzee habitat
ever been surveyed, especially

country, and the impact of the
Those areas
revisited."
gorilla
has a discon-
from
that have
its
former range
in
in
DRC has
the north of the
civil
war
is
unclear
gorilla
is
to
be
probably absent
the extreme west of DRC,
north of the Congo Riven'"'^
become
in
been surveyed need
The western lowland
and from the northwest
of
Biega NP, a density of 0.4/km' was reported," '"and

in
the Kokolopori
tinuous distribution east of the Lualaba River and
west
be widely distributed
parallel to the Albertine
forest."'"'"
The eastern lowland
to
in
Lac Tumba, and most recently
Uganda's
to
Congo River They
the north and east of the
appear
of
mountain
variants of the eastern
Lake Tanganyika.^' They also occur
There are currently conservation and
localities.
known
west, near the border with Congo.*^
bonobos estimated over
100 000.'"'" Bonobos are
all
more than
population
the last 10 years range from 10 000 to
hosts
of potential
has not been surveyed, however, which

to
DRC
Reserve" and are present
16.7al."
area occupied has been estimated at no
habitat
gorilla.
fragmented and
highly
discontinuous distribution (see
kml" Much
is
River.
The range
of
Bwindi Impenetrable NP, also includes part
Hence,
t.
population
largely restricted
is
as
from other eastern lowlands,
Bwindi
which
gorillas,
eastern
of
identified
not recognized as a subspecies.
distinctive
IP.
the Bas-Fleuve region, to
in
mouth
18 000
It
is
the
the western lowland gorilla \G.
and the central chimpanzee [Pan
the north of the
are
gorilla 16. beringei
small population
lowland gorillas, which has been
but
and eastern
Both the bonobo {Pan paniscus]
the northern sector of the
in
supports
distinctively different
in
2001' - about half the Virunga population.
It
extinct prior to 1980."
is
thought
to
have
The central chimp-
333
World Atlas
Map
16.7a
of Great Apes ano their Conservation
Bonobo and chimpanzee
distribution in the Democratic Republic of the
Congo
Data sources are provided at the end of ttiis

country profile
15E^
Central African
,-^'ii''
if
REPUBLIC
-1^^-r
V .,r-'
ti.--/
S-^
o o
BomuSNRO
SUDAN
Abumonbazi ^
^>.
FR
Lomako FR
'
"i
.,*='
tO
\
/'
/-
'^<,^
;-
.>-i;'-</o'
Bili-Uere
Garamba NP &
CAMEROON,
X^
.^
,-^.-.^^^-
WHS
Mondo Missa HZ
'f
/-,
.>?r.<^"
UGANDA
Albert
WHS
ard
'
.,
0"N
I
BURUNDI
Maniema FR
"'irp'/
/SBombo-
f V'^^
-^^"fey^*:
5SX
Lumene HZ
l4jKuru
&a
?
V
THE.CONCb"'""^"'
.Gf^,
REPUB.LIC OF
Matadi
,2î_
"V Luam^
DEMOCRATIC
Bosoaandja FaR
^
flEl
V TX,">BAS-
^^
<{
'y
KASAI OCCIDENTAL
(Proposed)
Mont Kabobo NR
^'*''\ (Proposed)
-
?,
'
Monts llombwe
NR
'
5S
fV:^--**:''
Kananga
Mbuji-Mayi
CABINDA
REPUBLIC
v'*^
ANGOLA
_Bushimaie
OF TANZANIA
Jy*V
HZX
Luama/
lac
,k,
Shaba HZ Upe,h^J'^J{^'^*'^
\
lO'S
Ml
O*
10^
LubudiV L-v
ANGOLA
Samppa
HR*
Lumumbashi
ZAMBIA
'1
IP?
Species
Estimated range
Bonobo observed 1996-2004

V-Vj- Central chimpanzee
iJiV.j^
Eastern chimpanzee presence alleged
^^-4' Nigeria -Cameroon chimpanzee
Bonobo observed
Bonobo observed before 1980
Bonobo presence alleged
Bonobo
Central chimpanzee observed
Nigeria-Cameroon chimpanzee observed
Nigeria-Cameroon chimpanzee observed before 1983
:-
after
1980
1940
1996-2003
15E
334
V-IJ" Bonobo
Eastern chimpanzee locally extinct since ca. 1940
Nigeria-Cameroon chimpanzee observed 1996-2003

after
1983
Eastern chimpanzee
Bili-Bondo ape
15^
anzee also formerly occurred

but there
the far west of DRC,
in
no infornnation on
is
present status
its
the Okapi
in
reserve during the conflict included the meat of at
chimpanzees. As figures
least three
or distribution.
bushmeat consumed
reserve." The
for the impact
of the conflict
on the chimpanzee population are not
THREATS
available,
urgent that fieldwork be carried out
Impacts of the 1996-2002 conflict
establish their conservation status.
apes
lations of all the great
decreasing.
7,
10. 26, i6,
53
have been badly affected towards the end

century. There
particularly
is
the highest
eastern
conflict,
human
little
densities
of
the 20th
DRC had some
war
1994 led
in
The Rwandan genocide and

an
to
influx of nearly
860 000 refugees were concentrated
NP
of the Virunga
(7
900 km'l.
the
civil
million
the vicinity
in
Some 332 000 had

NP,
Kahuzi-Biega
near
gathered
in
Rwanda and Burundi." About
from
refugees
of
Central Africa, with
in
000 km' originally established
area
an
of
DRCs
to protect
eastern lowland gorilla population."' ^ Large areas

of forest
for
were
cleared,'
'
fuelwood and food
protected areas."
1996,
In
in
into
in
DRC. By
were located within or
Many eastern lowland
NP and
late
Maiko
NP
be around
1998, there were estimated to
In
lowland gorillas
eastern
16 900
existence,'^ but following the war,
in
is difficult
it
to
know how many remain. Population surveys have

only been carried out
ists
in
few sites"" (see Status
in
Chapter
8),
with
many
conservation-
therefore being unwilling to provide an overall
estimate.
sector
is
It
no
certain that by 1999, the highland
NP had
percent) of Kahuzi-Biega
about 50 percent of
lost
gorilla population, including
its
88 percent of the gorillas habituated for tourism.'^
These habituated animals were

to shoot.
By 2001, the parks
estimated
to
be
000
in
particularly easy
gorilla
population,
995 census." had been
substantially reduced by hunting for
bushmeat."
spring 200A, park officials regained control of

of the park, but
(tin
ore),
In
much
estimated that 100 coltan, casserite
and gold mining areas were
still
used.''
Apart from a few well observed family groups, the
so there
is
need
a clear
for field
is
unknown,
surveys to help
establish their conservation status.
The Maiko NP
gorillas
victim to the
fell
to
situation for the eastern lowland gorilla
clearer
little
current status of the park's gorillas
war broke out
territory.
Kahuzi-Biega
incursions
to
"
civil
1999, all the national parks
near rebel
and the huge demand
led
is
is
and trends
DRC.^'
in
around 300 individuals per square kilometer

province of Kivu."
to
information about
chimpanzees
the current status of the
Before the
be
to
seems
lowland gorilla
The
popu-
gorilla,
DRC appear
of
Although no firm figures are
the eastern
available,
mountain
of the
With the exception
it
northern
limit
110
830 km') constitutes the
the eastern
of
lowland gorilla
in
large-scale slaughter of wildlife by fighters and
eastern DRC, with an estimated 859 individuals
refugees."" " The high price
reported present
of coltan
(columbium
and tantalum) ore between 1998 and 2000 led

further invasion of Kahuzi-Biega
NP and
to a
the Okapi
Faunal Reserve, by an estimated 10 000 and 3 000

people respectively.""
DRC
is
unusual
that there
in
are few fixed coltan mining concessions with secure
and so coltan
tenure,
is
generally mined with
in
1996, along with 4 000-5 000
chimpanzees." There were seven
gorilla
groups
here making up distinct northern, southern, and
extinct.'^ Gorillas are
thought
to
be
absent from large areas
of
western populations, but the
apparently suitable habitat
latter
in
is
the park." Threats
include hunting and mining, but there
human
is little
timber
shovels along rivers.' Professional hunters accom-
extraction and the
panied these miners and their dependents. Local
sparse." The impacts of coltan mining here are
warlords and rebel leaders are believed to have sold
unknown," so survey work
much
used
coltan
in
order to raise
money
for
hunting as well as warfare.""

the mid-1990s, between 12 800 and 21 900
30 530
were
estimated
to
live
Biega NP, and Okapi Faunal Reserve
of 7
in
113
in
eastern
726 km') supported a population
500-12 000 chimpanzees." Even prior
conflict,
issue,
the
km' area covered by the Maiko NP, Kahuzi-
DRC." Okapi
Gorilla
is
is
relatively
urgently needed.
Reserve (700 km') was set
up between the Maiko and Virunga NPs by two local
in
In
chimpanzees
The Tayna
weapons,
population
to the
commercial hunting and mining was an
with four large villages sited within
the
communities, the Batangi and Bamate Nations.

It
was
officially
reserve'
in
recognized by
DRC
as a 'private
2002, which simply indicates that
managed by
the
community rather than by the
it
is
state
wildlife
department. Preliminary surveys suggest
that
supports between 225 and 360 eastern
it
lowland gorillas. ''
'
Patrol
teams protect the
reserve wildlife from exploitation, and a 5
km
buffer
335

World Atlas
Map
the Democratic Republic of the Congo
in
20E
15E
25"E
this
country profile
3Q"E
>
(
.
^/^.
SUDAN
CENTRAL AFRICAN
RE PUBLIC
CAMEROON
Bomu SNR
(.
Gangala-na
oaramba MP
BodioHZ
&v(,HS
5"N
Mondo
Missa
HZ
Maika-
^
H
Penge
r-eafcv.
C.'.'
CSSko FR
>
(Proposed)
haut-
^,
Yangambi
L'l/'rj^.
...,
*:...,
>
, o-|
'Luo
^"^
"C*"
'^
Virunaa
KID sMîuc
// NP&WHS
Lservel^^ MOUNT
'ubandaka
jo,aan,
JOsangan,
WambaScR--^ KOKOLOPORI
^-
ong
Alhcrl
*
y>
TaJnaGorillart
MaikoNP>
Lake
FaR
.Okapi
J \&^NHS
FFR
UGANDA
'*""
'FR
npi
TSHIABERIMU
î-^
Lake Edward
Sarambwe
Gofflia
KASE
'^' ji-
BakanoFR
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rRWANDA
'
'
r- Kahua-Biega
'-'~{^^
Maniema^ "SKIVU _.^.V'
FRVy
DEMOCRATIC
Monts Itombwe
Kinshasa /"."Q^
'
"
REPUBLIC OF
/SBombo(_lLumene HZ
.''
5S
0\
(Proposed)
THE CONGO
iwonlKabobo
NR (Proposed)
'
GR
^
Matadi
Xananga
CABINDA
l_|-7
UNITED
^ Vepublic
I of'tanzania
MbujI-Mayi
Bushlmaie
Swa-KibulaV?
NP&WHS
'"
-BURUNDI
Luama'
'(5bas-fleuve
ANGOLA
NR
SR"
~Liike kivt
eLuama/
Shaba HZ
^ Kahemba
'-^
Lac
Upemtâ
Upemb
Tibaf^'
NP
P*r
''V(\
ID'S
Samppa
ANGOLA
Mountain
Western lowland
HR^
-Si
gorilla
gorilla
10"S
Lumumbashi
Species
Eastern lowland
MOUNTAINS
ZAMBIA
gorilla
Confirmed range
Eastern lowland
IS^S
f
Mountain
gorilla
15S
gorilla
Estimated range
U.ls Western lowland
''"'
-
336
ISÊ
750
500
km
gorilla
20E
25E
30'E
zone
protected from conversion. The Dian Fossey
is
Fund
Gorilla
Community Conser-
International's
Program
and
tielped to establish Tayna.
been largely unaffected by the war, but
access,
it
greatly
reduced.^'
Lomako
forest, for
other proposed reserves
There are
vicinity.
between 700 and
tentatively thought to be
gorillas
the
in
^00
Tayna and these adjacent areas. The
in
communities are worl<ing together through
eight
the Union of Associations for Gorilla Conservation
and Community Development
(UGADECI, and wish
to
DRC
Eastern
in
promote development
and bonobo
one northern part
bonobo
less
to
flight
Hunting Intensified
number
the
likely to
the
of
density;'" local
in
the late 1990s,
response
in
and an increase
in
commercial hunters entering the
of
The trade was limited by the war, but
now
return
"
accessible areas.'
is
that the conflict has subsided.
Bakumbule Primate Reserve,
include
Bakano Forest Reserve,
Reserve,
Ngira'Yitu
Community Reserve, and Punia
Reserve.
In
addition.
Sauvegarde de
In
example, nest counts indicated
to disruption of local agriculture
area.' "
Usala Gorilla
easier
reports suggest a combination of hunting losses
new
reserves
is
a 75 percent decrease in
through conservation rather than through unsustainable use of natural resources." The
those
in
density and
possible that populations have been
is
working with other local communities on seven
vation
human
areas with greater
Locale pour
Initiative
Nature (ILSN, Local
la
Safeguard Nature)
is
Gorilla
la
Initiative to
active over the Masisi territory,
and Action Communautaire pour
la
Protection
Protected areas less affected by
war
Salonga NP, which has two sections covering a

area of 36 560
km^
is
(and national park)

to
It
in Africa.
It
was created
in
1970
endemic species, including the bonobo.
protect
one
is
total
the largest rain-forest reserve
two protected areas supporting
of only
Mwenga lACPN-IM, Itombwe
the species (the other being the Luo Reserve for
Mwenga Community
Action for the Protection of
Scientific
Nature!
the
de
Nature Itombwe
la
is
involved
was thought
forest
gorillas
in
in
Itombwe area." Itombwe
have supported about
to
155
1998," and to have been under moderate
hunting pressure at that time." The forest area
proposed nature reserves (Monts
includes two
Itombwe and Mont Kabobol,

(Maniemal, and a
its
location
the center of the country has
DRC. The Mabali
sites in
like
Luo,
meant
it
is
Scientific
Reserve (19 km'),
center for bonobo research, and
receives local government protection.'"
Other protected areas that host great apes
Bososandja Faunal Reserve [34 km'), and Rubi-Tele
It
now occupies
only about 13 percent

of
Hunting Reserve
Forest Reserve
the fragmentation of the populations involved.'" The
WWF-The
small Masisi and Mount Tshiaberimu populations
1980s,
are particularly vulnerable.''
animals
boundary
In
1998,
lived in the forests
Tshiaberimu'^ and 28
refugee
in
is
Salonga NP, and
in
has been less affected by the conflicts than other
reserve iLuamal.
geographic range, which reflects the extent
tensive
358 km'|.' There
probably the highest of any of the gorilla
is
that 16
at
include the Bili-Uere Hunting Zone
subspecies."
of its
reserve
forest
Wamba,
rate of habitat loss for the eastern lowland
The
gorilla
game
Research,
as yet no reported logging threat
in
was thought
around Mount
it
the Masisi area."
On the
Mount Tshiaberimu, there was
of
agricultural
encroachment during the

gorillas,
or about 5 percent of the population, were killed as

a direct result of military activity
along with an
unknown number
Although an end
forests of
DRC
and retreats
from
to the
of
992
to 2000,
chimpanzees.^'
war has been declared,
for rebel forces, leading to continuing
largely inaccessible by park staff,
the gorillas
the
continue to serve as hiding places
hunting," The lowland areas of the parks are
is
being set aside by the logging
approval
for
rupted by the
''
in
the
company
The process
national designation
was
of
inter-
war
Threats
in
peace time
The
relative
the
country's
mean
that,
remoteness
of
prime timber areas and
poor transportation infrastructure
until
now, only low-volume, selective
logging has been profitable, and then only
in
limited
areas along large rivers. This has effectively protected

5 320
much
km'
cleared
in
of
the forest of DRC.
of forest (about
OA
It
is
estimated that
percent of the
the country each year"
total! is
Forest Code
was established in January 2003, under which the

state owns all forests and defines legitimate uses
for them. Legal mechanisms and a zoning system
have
are to follow. Concerns have been raised that there
unclear"
to
suggested by
first
Global Consen'ation Organization
of
still
and the status
Most bonobo populations are thought
080 km'l. The proposed Lomako
600 km'l was
Siforzal that held the concession.^

its
000 km'),
ex-
crisis.''
Between 12 and 17 Virunga mountain
(9
(3
(6
337
World Atlas
of Great Apes ano their Conservation
and around the mountain
in
Hunting
of this
20 years due
protected
mammals, such
more serious
National Park with
in
elephant and gorilla
the
as antelopes, had represented the
By 998,
threat.
at least
montane sector
to
catch other
to
each habituated eastern lowland
hand
from being
gorillas
Wire snares set
food.
Staff at Kahuzi-Biega
Free Foundahon
the last
Congo Basin also
the east
in
many eastern lowland
for
killed
Redmond/UNESCO/Born
in
dedicated protection and support
to
taboos
Historically,
Ian
rare
governments and conservation groups."
from
skulls.
range area.
gorilla
become
species has
'^
snares."
Kahuzi-Biega
of
one individual
group
gorilla
NP
had
As noted above, by 2001, most
these gorillas had been
in
lost a
of
killed.
are no plans to recognize the resource rights of
people traditionally resident

there has been
little civil
in
the forest, and that
society involvennent
in
the
The Ministry
low (US$0.06/hal, so large areas must be
and Tourism
licensed to timber companies for government to

obtain
income from
significant
source. The
this
World Bank expects that 600 000 km*
be zoned
will
as production forests.^'"
Timber extraction
will
made
be
easier by
of
donor funds promised
April 2004." " This
in
equivalent to the reconstruction of about

of the country's 2
km
of
its
800
km
of
080
is
km
paved roads or 5 400
unpaved roads." As trade routes are
revitalized, there
is
danger that the bushmeat
trade will follow the timber trade.
existed for all ape species
in
The
and movement
loss of these taboos,
without
them
into
bonobos
of
DRC.
is
example, local people near
have traditionally refrained from hunting

^^
However,
the government body responsible
The Nature Conservation
parks. The 1982 hunting act (Law 82.0021 defines
reserves
animals
prohibited.
game
and
which
for
Unusually,
permitted
with
in
in
parts of
others."
bonobos have also been hunted
Historically,
their
many
'"
for
supposed medicinal and/or magical value,
charms made from body parts being
some
areas." Relatively few people
around most
of the
so hunting pressure
range
is
of the
available
live in
and
bonobo, however,
low overall, though bonobos
are very vulnerable to wire snares set to catch other

a widespread practice,
and one
in
which a very few people can set a large number
of
species. This
traps, all of
is
which remain a threat
for years
even
if
they are abandoned or lost by the snare-setter
A taboo
against eating gorilla meat
still
was
was
restored following a campaign
by the Lukuru Wildlife Research Project team.^""
International
DRC
agreements
ratified
Biological
or acceded
Diversity
in
exists
in
on Migratory Species
to
Combat
to
1994,
the Convention
the
in
in
in
Convention
Sites:
in
in
UN
1997,

Natural Resources
of
Wild
1976, the Convention
1990, the
Desertification
on
Convention on
Endangered Species
International Trade
in
Zone Dekese
There were no such incidents, and their
protected status
Fauna and Flora (CITES)
avoided
In
1997 with written permission for each
in
for
is
own
(Province Kasai Occidental!, bonobo hunting
the
DRC, and regarded as a delicacy
are
regional
1985,
right to set their
hunting seasons, bypassing this law.
in
meat." Bonobo meat
lists
trapping
regulations on species protection and to specify
mid-1980s hunters began taking bonobos
for religious reasons."'
and
since
governments have had the
and
reserves
hunting
people
great ape range areas,
significant threat. For
Wamba
different areas of
Environment, Nature Conservation,
is
Act of 1969 (Ordinance-Law 69.0411 defines national
incident.
Although not universal, hunting taboos have
of
for nature conservation.
faunal
planned new roads, supported by US$270 million
338
formulation of the code.'" Forest exploitation taxes

are
LEGISLATION
of
Convention
the African
Nature and
1976, and the World Heritage
1974. There are five World Heritage
Garamba NP, Kahuzi-Biega NP, Salonga NP,
Okapi Faunal Reserve, and Virunga NP.

as World Heritage Sites
pressures.
DRC
in
All are listed
Danger due
also participates
in
to
human
UNESCO's Man
and Biosphere (MAB) Programme.
Protected areas
There are four
mam
categories of protected area
DRC: national parks
(nine sites],
game
in
reserves
lone
site],
and faunal
"'^ Other designations include
forest reserves (seven sites],
reserves Itwo
sites!.
200A, bonobo conservation and research
In
activities -
underway and proposed by the Centre
areas set aside for scientific research, or as hunting
de Recherche en Ecologie
zones and nature reserves. The existing protected
Wildlife
195^26 km-
area network covers an estimated

(8
Management of
and game reserves
percent of the land areal.'^
national parks, faunal reserves,

is
delegated
the
to
Conservation de
ages
Congolais pour
in
la
Nature (ICCN), which also man-
la
scientific research. Effective control of
protected areas
in
Institut
many
the east of the country has been
the hands of rebel authorities
in
recent years.
Sauvage,
Wamba Committee
WCS, and
Most research
in
in
DRC has taken
and
place
the east and north of the country. Research sites
include Kahuzi-Biega NP.'-' Salonga NP," Okapi
Faunal Reserve,"
Luo Reserve
in
Lomako
Wamba
Scientific Reserve, the
Research," and Lui Kotal
for Scientific
Research was scaled down
forest."
during the war, but persisted at several

In
particular, the
Max Planck
Lukuru
Wildlife
Institute, the
Natural Sciences
in
DRC
included biomonitoring, building community relations,
economic development
the
Research Project,
Center
Research for
of
ICRSNl, and the Wildlife
war.
civil
CRSN
to
work
researchers
continued a research project on sympatric gorillas
and chimpanzees
in
Kahuzi-Biega NP,
in
cooper-
National organizations active
pour
DRC
in
la
in
ape conser-
include the Association de
Conservation et
le
programs, infrastructure building and rehabilisurveying and mapping, training
tation, inventory,
of
poaching patrols, and research.^'
center, extending national park boundaries, formalizing
the awareness-raising programs, demarcat-
ing protected area boundaries, building
awareness
among logging concessionaires, developing land

management plans, providing appropriate equipment, training, and support
the
ICCN, and revitalizing
to
Lomami-Lualaba conservation
site.
Ecotourism
Gorilla tourism brought 2
NP between
800
visitors to the
1986 and 1990,
Virunga
who between them
spent an estimated US$250 000. Meanwhile, be-
tween 1988 and 1991, 2 000

Biega
NP
visitors to the
Kahuzi-
generated an estimated US$200 000."
Ecotourism income has been minimal
Femmes
through poaching
Kahuzi-Biega
NP
of
is
for the past
habituated gorilla groups

a
in
the
major setback.
Developpement Durable
Sanctuaries and rehabilitation
Programme
The Lola ya Bonobo sanctuary
pement Economique du
more people
conservation and research, awareness-raising
in
(AFECOD), the Pole Pole Foundation IPOPOF], and

d'Appui aux
micro-projects
decade, however, due to the conflicts; the death
ation with Kyoto University'"
vation
of
outside the Salonga NP, employing
field sites.'"
Conservation Society (WCSI continued
throughout
Bonobo Research,
Other goals included establishing an education
field projects
into the populations, ecology,
socialbehaviorof great apes
for
Lukuru
Institute, Vie
the Zoological Society of Milwaukee -
and support
Conservation and
Forestrie,
et
Research Project, Max Planck
de Develop-
Initiatives
Kivu iPAIDEKl. Typically,
is
located 25
km
for
orphan bonobos
from Kinshasa, and run by Les
these focus on community projects and alternative

Jo Thompson/Lukuru Wildlife Research Project
livelihoods. Active
international nongovernmental
organizations include the International Gorilla Conservation
Programme
IVirunga NP),
WWF
(Virunga
Wild bonobo
in
Bososandja Community
Forest.
and Garamba NPsl, Lukuru Wilalife Research

Project
(Salonga
NP|,
WCS
Faunal Reserve, and the wider
(Salonga
Ituri
NP, Okapi
Forest Reserve),
the Zoological Society of Milwaukee (Salonga NP),
and the Dian Fossey
Gorilla
Fund (DFGFl (Mount
Tshiaberimu and Virunga Conservation Center). For

example,
DFGF aims
to
establisn
conservation zone between Maiko
community
NP and
Kahuzi-
Biega NP.'^ Unusually, the Lukuru Wildlife Research

Project has purchased land rights to the Boso-
sandja
range
Faunal Reserve (which incorporates the

of the
Bososandja bonobo community).
339
World Atlas
holds important populations of eastern lowland gorillas and chimpanzees;
NP
reclaim the parts of Kahuzi-Biega
still
outside the rangers' control and survey the
how many apes have
area to establish
vived
sur-
onslaught of bushmeat hunters
the
feeding the coltan miners;
develop community conservation

create jobs
areas
in
of rural
initiatives to
poverty
tions important for great apes,
loca-
in
such as around
the village of Lomako;
The
at
gorilla
census team
strengthen existing laws protecting great apes
and improve awareness among law enforce-
Virunga National
ment agencies and the
Parl<.
Martha
courts;
Robbins
increase resources for sanctuaries to care
Amis des Bonobos du Congo.

38 bonobos, with numbers
May
In
2004,
it
had
for confiscated
infants arriving at
of
potential
the center increasing since the end of the war.
infant
apes and develop the
sanctuaries for conservation
of
education.
Investigations for an appropriate site for eventual
reintroductions are underway.^'
an ICCN meeting decided

the
Goma
In
November
to create a
region for confiscated
2004,
sanctuary
infant
in
eastern
Lowland gorillas."
An
action plan for
and
endorsed
sited close to
calls for regional surveys to
bonobo populations, conservation education
throughout DRC, economic benefits for local people
Kahuzi-Biega NP, cared for 13 orphaned chimp-
from conservation projects, and enforcement
anzees and one bonobo
antipoaching and habitat protection laws.
sanctuary
is
at the
end
under the management
of 2003.
of
The
ICCN, and
2003, an international
In
supported by the German overseas development
Japan sought
agency GTZ, the Born Free Foundation, the
national Primate
Jane Goodall
League
Protection
Institute (JGIl,
Inter-
(IPPLl,
the
and the Pan African
Sanctuary Alliance IPASA).'
number
the
to build
of
recommendations
bonobo.
These included
of
research
facilities lat
Lomako
The National Biodiversity Strategy and Action Plan
development
need
for the conservation of gorillas
and bonobos, which are highlighted within one

the priority projects - Project
rehabilitation
3:
of
Protection and
plans for threatened species and
2002
to
for conservation of
field
surveys
held
in
Kinshasa
discuss conservation action for great
(at
Kokolopori, Lac Tumba, and
forest
and
management plans (for

NP), community
Salonga
activities
(at
Wamba
and Lukuru/
Kotall,
[at
[at Lui
and continuing or establishing research
Wamba,
Salonga, Lukuru/Bososandja, and Lui
Kotall.
Various recommendations have been
A GRASP workshop was
in
made
It
Bososandjal, extending reserve boundaries
ecosystems."
in
plan.'"
of
Kokolopori, Lac Tumba, and Lomakol, construction

identifies the
workshop held
on the 1995
Lui Kotall, completion of
action
in
specific protected areas to
apes. Prior to the coltan rush,
it
made
for
conserve great
was suggested
that
DRC, from which a draft action plan was

produced.'" " The most urgent recommendations
conservation efforts for eastern lowland gorillas
from the working groups
Kahuzi-Biega
apes
in
at the
meeting were:
should be focused on the lowland sector
It
undertake surveys
of
little-known areas to
establish which apes survive and where,

particular
which
is
in
the
Mayombe
forest,
contiguous with forests
in
Bas-Fleuve,
in
Angola's
Cabinda province;
rehabilitate the
360
1995"
in
lUCN Species Survival
the
Commission. The plan

identify
The Lwiro Primate Sanctuary,
bonobos was published
by
neglected Maiko NP, which
NP and
had been considered essential
forest corridor
populations
to
to
maintain the
between the lowland and montane

ensure gene flow between them.^'"
Ivlore recently, calls
have been
made
for crisis
agement and damage assessment and

The Itombwe
of the
the adjacent Kasese region.^*
forest
man-
limitation.
has also been recommended
as a focus for the conservation of eastern lowland
and
gorillas,'^
population
its
important given the loss of so
seems even more

many Kahuzi-Biega
bonobos, the designation
Lualaba conser^âtion
gorillas.
Overall
purchase
bonobos include the
for
priorities
concessions
of forestry
these
sites,
as well as
as nationally protected
site,
Lomako
areas, and the creation of the proposed
Forest Reserve."
key areas for
in
of
Bososandja Community Forest and the Lomami-
'
FURTHER READING
Hart, J.A., Hall, J.S. (19961 Status of Eastern Zaire's forest parks and reserves. Conservation Biology ^0
Hicks, T.H. 120051 The
Bondo Mystery Apes, Winter 200i
121:
316-324.
Field Data. Unpublished report, http://www.karlammann.
corn/bondc-winter200i.html. Accessed June 8 2005.

Rainforest Foundation 12004)
New
Threats to the Forests and Forest Peoples of the Democratic Republic of Congo.
Paper February 2004. The Rainforest Foundation, London, http://vmw.ra1nforestfoundat10nuk.org/f1le5/
Briefing
RF%20UK%20Briefing%20Paper%20on%20ORC%20Februar/%2004.pdf Accessed June

RednDond,
(2001) Coltan
I.
Boom.
Gorilla Bust. Report for the Dian Fossey Gorilla
2 2004.
Fund (Europe) and Born Free
Foundation. http://vAww. bornfree.org.uk/coltan.
Thompson,
Hohmann,
J.,
G., Furuichi, T.,
eds (2003) Bonobo Workshop: Behaviour, Ecology and Conservation of Wild
Bonobos. inuyama, Japan.
UNSC
(2002) Final Report of the Panel of Experts on the Illegal Exploitation of Natural Resources
Wealth
the
of
Democratic Republic of the
Congo.
United
Nations
Security
and Other Forms

New/
Council,
of
York.
http://www.natural-re5ources.org/nninerals/CD/docs/other/N0262179.pdf. Accessed June 18 2004.
MAP DATA SOURCES

and b Great apes data are based on the following source:
liâps 16.7a
Butynski, T.M. (2001) Africa's great apes,
Stevens,
E.F.,
Arluke,
A.,
in:
eds. Great Apes
T.S.,
Hutchins, M.,
Ivlaple, T.L.,
Norton,
and Humans: The Ethics of Coexistence. Sm\lUsoman
B.,
Rowan,
A.,
Institution Press,

With additional data by personal communication from Blake,
(20031, Plumptre, A. 12004), Ron,
Hall, J.S., Saltonstall, K.
Grauer's
Hohmann,
Inogwabini,
(2003) Culture
Cultural Anthropology AA
Hohmann,
Ilambu,
News
Grossman,
0.,
Report
in
J.,
to
Omari,
F.,
12003),
Lanjouw,
A.
and Gray, M.
abundance and conservation status
of
bonobos? Between-species and within-species
in
variation
in
behaviour
563-571.
Kotal - a
new
site for field
Mbenzo,
P.,
research on bonobos
in
the Salonga National Park.
Blake, S. (2005) Monitoring of the Illegal Killing of Elephants
and Ape Population Surveys
CITES MIKE and the government
Williamson,
(19981 Distribution,
I.
J.
Pan
2b-27
i2\:
Forests: Elephant
Kalpers,
14):
G., Fruth, B. (2003) Lui
Africa
B-l.,
Thompson,
122-130.
gorilla. Oryx 32:
G., Fruth. B.
S. (2005), Furuichi, T. 12003),
[2003], Stokes, E. 12003],
T.
E.A.,
in
of the
in
Central African
the Salonga National Park, Democratic Republic of Congo.
Democratic Republic
A.,
of the
Nzamurambaho.
Congo.
A., Lola, N.,
Mugiri. G. (2003) Gorillas
the crossfire: population dynamics of the Virunga mountain gorillas over the past three decades. Oryx 37:
326-337.
Thompson,
J.
(2005) Field research at Lukuru, Democratic Republic of Congo.
Wamba Committee
for
Bonobo Research
Pan
Africa
News
10
(21:
21-22.
(2004) Latest News. http://vAA/.pri. kyoto-u.ac.jp/shakai-seitai/shakai/
B0NOBOHP/English/News_e.html. Accessed May
2005.
ACKNOWLEDGMENTS
Many thanks
Yamagiwa
to
reviewers Jo
Thompson ILukuru
(Kyoto University) for their valuable
Wildlife
comments on
Research
Project],
Annette Lanjouw IIGCP), and Juichi
AUTHOR
Nigel Varty,
UNEP World
341
World Atlas
REPUBLIC OF EQUATORIAL GUINEA

Brigid Barry

The Republic
of
pressure on Equatorial Guinea's wild animals,
Equatorial Guinea
smallest countries
Africa.
in
one
is
of the
two principal
Its
including the great apes, as a source of
meat
for the
growing affluent population.
regions are Rio Muni, bordering the Bight of Biafra
between Gabon and Cameroon, and the island
The
Bioko.
26 017 km'
climate
equatorial with a
is
of
The
km'', of
of
which
mean temperature
of
17-34C1. The average annual
Muni
precipitation in Rio
southern Bioko
in
28 051
is
the continental Rio Muni. The
in
is
26C (range
while
area
total land
it
is
less than 4 000
can reach
mm,
000 mm.''
11
troglodytes] are both present
The current numbers

early
1
1980s,
of
in
mainland Rio Muni.
apes are unclear
was estimated
it
the
In
that there
were
000-2 000 gorillas and 600-1 500 chimpanzees,
with 150-500 breeding female chimpanzees.''''*
Fang, and the main ethnic groups are Fang, Bubi,
1989-1990 census concluded that depending upon
Annobonese, Ndowe, Kombe, and Bujebas.^'
the calculation applied, there could be 990-2 450
independence from Spain was
ratified in
1968
and Equatorial Guinea remains the only Spanishspeaking country
Under
1979.
Equatorial Guinea
sub-Saharan
in
is
economic
behind Angola and
Africa,
US$2.2
billion
[2002 estimate),' largely as a
result of the expansion in the
GDP
in
increased by 71
.2
2001, and around
makes
percent
and gas sector'
1997, 45.5 percent
in
15 percent
Guinea
Equatorial
oil
dramatically, up from 358 000
The population
is
in
2003, which
in
world's
the
growing economy' The population
2004.'
changes.
The country's gross domestic product
Nigeria.
IGDP)
power since
in
has experienced
the third greatest producer of
is
is
fastest
also rising
1991 to 523 051
growing
in
around 2,5
at
percent per year according to 2000 estimates.'

infrastructure,
health,
sanitation,
and education
remain underdeveloped.
apes
in
of the
most people
Jones and Sabater Pi" spent 18 months

1967
still
popula-
work
in
Aninzok, and Monte Okoro Biko Inow called
Monte
Mitral.
The conservation status
was considered
species
of
two
to
cassava, sugar cane, maize,
pineapples, bananas, and plantain."
evidence
of
livestock
and chickens. The lack
attributed
"" There
repeated failed attempts
to
the tsetse
fly
of
to
is
breed
success
is
(vector of trypano-
in
stable groups
home ranges
were found
the ground
in
open areas, with nests close

thick vegetation. Central
in
anzees were encountered

1.53/km'
23 individuals. Their
mostly
remains
undeveloped
home ranges averaged
the upper strata of the forests, with
in
in
the canopy of small trees surrounded
by taller primary trees.
Gonzalez-Kirchner'" studied western lowland
in
385
this,
km
of
gorillas per
throughout Rio Muni for 18
to 1990). Gorillas
783
gorilla
km
were present
of transects.
Based on
density of 0.45 of nesting
square kilometer was estimated
for the surveyed areas. Site densities
from 0.12/km'
southeast
also
at densities of 0.31-
approximately 15 km'. They were observed
0.71/km'
Fishing
to
chimp-
fission-fusion groups of two to
in
European
respectively
human
densities
averaged approximately 6.75 km' and they
somiasis] and the low prices of imported, frozen
chickens
of both
to
12 individuals. Their
months (1989
(cacahuetel,
due
gorilla
varied from 0.58 to 0.86/km'
malanga {Xanthosoma
nuts
critical
Western lowland
activities.
gorilla populations
sweet potatoes, pea-
in
1968 at Monte Alen, Abuminzok-
to
the agricultural sector^* The main food crops are

spp.l,
on the density
Equatorial Guinea are known.
nests
With an estimated 64 percent

tion living in rural areas,
individuals.'" To date only four studies

of
Africa. President
his rule, the country
demographic and
drastic
oil
sub-Saharan
in
Nguema Mbasogo has been
Obiang
in
in
the Rio
Campo
ranged
basin to
the Nsork highlands region
in
the
of the country. Gorilla densities at
Rio Muni, with
Evinayong (0.26/km') and Niefang l0.59/km')
European and Nigerian trawlers exploiting the
were considerably lower than those estimated
waters."'
342
gorilla {GoriUa gorilla gorilla]
and the central chimpanzee [Pan troglodytes
"
languages are Spanish, French, and
official

The western lowland
relatively
^'
in
Limited domestic protein sources put
in
the
1960s at Evinayong 10.58/km') and
AFRICA: Equatorial Guinea
Map
in
Equatorial Guinea
th.s
country profile
CD
O i
'i^
(D
,p
ra
f>
r-
r-
ni
CI)
(1)
(1)
tl)
Si
cn
m
n
t
fc
CD
(D
CD
P
LL
M
(0
t
r
CO
(Q
p
CO
o>
ti
%
r
CO
a3
t;
CO
h s a 1 ^
(D
?,
T3
F
a
(0
Ul
lP
*-.-^-.;i;:fc=t.tl.k-=t
u
^
._
<
U
X.
*l
1
3
EQUATORI ^
Z;
--
3^3
World Atlas
Ghiurghi and Puit" spent four months [2003 to

200AI recording gorilla and chimpanzee nests
km
along 68.3
of transects
the southern
in
extension of the Monte Alen NP. The nests of
were
the two species together
at a
low density,
0.82 nests per kilometer or 24.18 nests per
square kilometer There was very
dence
areas
presence
of a gorilla
the park, which
of
absence
of
little
evi-
the southern
in
may result from the

[Afromomum and
key food plants
Marantaceae speciesl.
THREATS
Bushmeat hunting
The greatest current threat
15
bushmeat
hunting.
to
Muni
Rio
in
meat as an essential
largest ethnic group, regard
component
apes
The Fang, representing the
of their diet.''
Fresh meat and
fish is
preferred over frozen alternatives. While domestic
meat options
do
hunter with a dead
and
mandrill, Equatorial
of
Guinea.
Noelle Kumpc-I
Niefang
A subsidiary study
l0.86/km^l."
reported an abundant population of central
chimpanzees
same
highlands
the Rio
in
Presence
time.''
Campo
area
of gorillas in the
was confirmed
same
1994 as part
studied the distribution and
of a
longer primate study
Monte Alen National Park

1997 extension
They
south.
(NP), prior to
in
its
to include tônte l^litra in the
identified
208 gorilla
Gorillas
were most abundant
forests
(5.15
in
nests.
secondary
nests per kilometer of the
surveyed area),
individuals.
and a source
in
mean group
Most were found
in
sizes of 3.3
the
montane
areas crossed by the Niefang-Bicurga road,
Wele
Rivers.
for a large
all
of the
Lana-
Chimpanzees were found
in all
In
bushmeat trapping and
people from Sendje, a village
actively shooting or trapping,
were counted both

Box
13.4). In a
inside
mean group
size of 2.3 individuals. This
evidence of ape hunting,
but
concluded that they were not threatened

within the park.
of
5 months, 80
400 people on
and nearly 5 000 traps
made
previous study
over a period of
11376 kg
of
bushmeat. This was declared an
unsustainable rate."' " Hunters, though Increasing

in
abundance, are currently fewer than trappers
because guns and cartridges are expensive and not

readily available
in
The apes are
Equatorial Guinea."
able, often collectively, to
from wire snares, although
escape
and/or limb loss
Injury
is
possible. However, hunters are increasingly killing
bushmeat, and as guns become more
apes become easier
nests per kilometer of the surveyed area],
of
16 months, the village of Sendje alone harvested
for
(5.35
hunting.'
Wele River
and outside the park'* (see
able,
study found
of the
the edge of the Monte Alen NP, were recorded as
apes
open primary forest
of rural
have experienced more extreme hunting pressure
areas surveyed, with 333 nests detected. They

in
number
the rural villages studied throughout Rio
were abundant
with a
3a
Income
of
The regions north and south
with fewer observations around the low-lying
Lana River and the confluence
that bush-
people. With the exception of coastal towns like
Muni are active
population densities of apes for three months

in
means
than others. ' During a study period
Mba"
with the Fang
combination with a scarcity
both a substantial part of the people's diet
Cogo,
area.'"
Garcia and
IS
Nsork
the
more popular
other fresh meat sources, this
meat
at the
in
is
slightly cheaper.' In
by a 1998 survey,
which also found chimpanzee nests
imported from Cameroon!
(e.g. cattle
bushmeat
exist,
targets.
Among
the Fang subtribes, apes, especially gorillas,

traditionally believed to
cause
Infertility In
avail-
several of
were
women
if
demand and
the cost of transporting the heavy corpses was not
worth the market return. However, many previously
eaten.' This
meant
that there
was
little
AFRICA: Equatorial Guinea
taboo species are becoming marketable.

ago,
A decade
'^
bushmeat was mostly consumed
locally,''''
meat
but the improving infrastructure allows
be
to
The two main bushmeat markets in Rio Muni

in Bata; the Central and Mondoasi
are both located
Markets. Tons
of
all
little is
done
to
main
on the two roads entering Bata,
few African countries that
of
meat are
No
Convention.
of forest
was thought
the whole of Equatorial Guinea
with an average annual decrease of
between 1990 and 2000
of
the area
110
of
km^
forest cover
or 0.6 percent."
plantations were
for
available.
Prior to the 1960s, large areas
forest of Rio
to
2000,
in
Statistics
not
of the
Muni were destroyed or degraded by
commercial agriculture and logging. Following

periods of political and social unrest led
abandonment
to
this,
the
commercial logging schemes,
of
emigration, and concentration of the

lation in the coastal regions.
The
human popu-
political stability
under President Obiang's rule has allowed an

increase
in
forest exploitation
and from the mid-
1980s on there was a further estimated 20 percent

reduction
In
in
in
In
of
the
Cameroon. The meeting
in
new
create
to
and poaching'".
of
2000, the
In
number
timber was logged
logging
Protected Areas
was passed, which increased
Equatorial Guinea
the
Law on
protected
illegal
areas
of protected
covering a total
to 13,
area of 5 860 km^ (approximately 20 percent of the

country). Ten are found
in
Rio Muni: two national
parks (Los Altos de Nsork, Monte Alenl;
five
natural
reserves (Estuano del Rio Muni, Rio Campo, Monte
Temelon, Punta Llende, Corisco y Elobeyesl; one

reserve (Playa Nendyi); and two natural
scientific
monuments
Alen
is
NP
IPiedra Bere, Piedra Nzasl."
covers an area
of
dense
" Monte
and
tropical forest,
the largest and most effectively protected of these
areas.''
The park has been
past and so
largely
is
selectively logged in the
made up
secondary
of
Hunting, logging, and agriculture are
forest.
officially
pro-
000 km" park, but there are
logging concessions running up to
boundaries.'
its
Rio Muni for export. Logging concessions have
been sold over most
of the country, including large
parts of the protected areas."^'

in
1999, Equatorial Guinea participated
Yaounde Forest Summit
hibited within the 2
forest cover'^
1999 alone, 788 000 m'
Man and
established the Yaounde Forest Declaration, which
rare.
Habitat destruction
in
the
of
Biosphere (MABI Programme."'
areas and to have plans to combat
Approximately 17 520 km^
one
have been designated as
sites
committed participants
remain
is
It
not part of the 1968
is
Biosphere Reserves under UNESCO's
monitor or prevent any bushmeat
and seizures
traders,
on Biological Diversity (19941.
and Natural Resources or the World Heritage
passing through. Bribes are extracted from bush-
meat
of Wild
Endangered Species
in
African Convention on the Conservation of Nature
positioned at the
guards are
forest
military barriers
a signatory to the Convention on
the
to
traders and bush taxis, and openly traded. Although

state
is
International Trade
over the country by
bushmeat are brought
Bata markets annually from
Equatorial Guinea
Fauna and Flora ICITESI 119921 and the Convention
markets/'
distributed to the larger city
the
In
particular, forests
Micomeseng-Ebebiyin-Mongomo region
in
Conservation and
is
European Union
part of the
program. Conservation and Rational Use
Ecosystems
the northeast have been severely damaged.
field projects
Equatorial Guinea
started
in
in
1992.
The
role of
of
Forest
(ECOFAC), which
Central Africa
ECOFAC
is
to
support the
Monte Alen NP, and
has
The pet trade
management
The growing economy has attracted non-African
trained park rangers from villages surrounding the
immigrants
to
the
Rio
expatriates regularly buy
Muni region. Individual

live
wild animals as pets,
mainly African grey parrots [Psittacus erithacus).
Although
illegal,
apes are seen as

that
this
'cute'
trade
is
Baby
and some expatriates believe
purchasing them saves them from being eaten
by locals. Hunters are reported to
apes
increasing.
to
kill
entire troops of
capture one orphan for sale.' They can earn
more by selling one baby chimpanzee to an expatriate than selling 20 blue duikers [Cephabphus
monticola] to the
bushmeat
market.''^'
of the
park to police
monitor flagship
activities,
illegal
it
species such as the apes, and to guide visiting
researchers and ecotourists.

structed a guesthouse
in
ECOFAC has
con-
the park to encourage
community-based conservation through tourism

revenues.'" Limited resources and a gap
combined with lack
of
tourism potential of the

trapping
still
success
site,
in
mean
in
funds,
developing the
that hunting
and
continue within the park. The park's
great apes are
seldom
killed this way."'^^
Nationally, conservation issues are adminis-
365
World Atlas
tered by the Institute Nacional de Desarrollo Forestal
Gestidn del Sistema de Areas Protegidas
(INDEFORl, a conservation body within the Ministry

of Infrastructure
and Forests IMinisterio de
estructuras y Bosques). Most of the
which places
botanists,
management programs
limits
Infra-
field staff
are
on the research and
set up for
apes and other

Research
A longer study on
is
needed
the ape populations of Rio Muni
reassess the densities and areas of
to
high ape conservation priority Research
in
has been sporadic and the information
now
is
the past
out of
date. Possible research projects are outlined here.
vertebrates.
In
2002, a
team from the Zoological Society
London and Imperial College London started

ject
on the sustainability
Muni,
in
bushmeat hunting
of
of
in
Rio
Initial
the
of
populations,
conjunction with ECOFAC, INDEFOR, and
Conservation International.
a study using nests as a proxy for the relative
In
abundance
a pro-
apparent differences
work focused on
and chimpanzee
in
that there
were
the distributions of
the two species between
open and dense
the incentives for hunting for villagers living close to
primary
Monte Alen NP, asking what socioeconomic circum-
studies might elucidate whether these
stances drive the decision to hunt, prey selection,
ferences result from species-specific habitat
and impacts on the
selection
viability of
Urban demand and the
prey populations."
role of
ences within the regional capital

analyzed.' Further
work
bushmeat
of livelihoods
in
terms
consumer
of
Bata were also
bushmeat
The Durrell
carried out another
with
INDEFOR
bushmeat study
for six
and
of agri-
tion study carried out by
if
confirmed
Gonzalez-Kirchners study'" found high den-
area,*
limited, at least by the early
it
bushmeat from an economic
general,
In
yields.
Despite the presence of orphan chimpanzees, there
to
are no primate sanctuaries or rehabilitation centers

Equatorial Guinea.
of
but one
(a
Bata found
private institution)
60 schools.^
hunting should be controlled inside
Ecotourism could use Equatorial Guinea's
population. Unfortunately,
in
spite of existing infra-
structure at Monte Alen and growing facilities
in
Bata, the large military presence has ensured that
international tourism has

find
to travel
it
difficult to
As hunting
is
managed
thought
ape populations
hibiting legislation
in
porting,
ing
in
to within sustainable
be the biggest threat
Equatorial Guinea, the pro-
needs
and trading
to
to
in
be enforced and stricter

place for
killing,
trans-
great apes. Improved train-
and management would help forest guards and
park rangers to carry out their part
in this
work.
Conservation organizations and government agen-
natural beauty to provide an income for the local
may
A
to
as an actively protected area.'^
controls and fines put
that environmental education, including conservain all
990s.
Hunting
Sanctuaries, education, and ecotourism
was absent
would help
of this district
demand
protected areas and
tion,
in this
which indicates that hunting impacts
had been
in
and ecological perspective.'
in
Nsork highlands. High
elephant densities were also recorded
to
bushmeat consump-
early 2004
by
sities of gorillas in the
John Fa and colleagues
in
management
conjunction
establish
A survey conducted
be high, the area should
to
Unit, Oxford
in
country with the aim of understanding the
in
of
be assessed,
to
ECOFAC.
and market data were gathered throughout the
of
past hunting
1989-1990. The current status
thorough study
and consumption
dif-
apparently had high chimpanzee
in
the 1990s. Household interviews, hunter interviews,
for
Further
forest.
to
be included within protective
hunting.'"
months from October 2003
March 200^. This followed up
related
chimpanzee densities needs
Wildlife Conservation Trust, Jersey
and the Wildlife Conservation Research
and secondary
or are
Campo
Rio
densities
and food security
across Rio Muni,^ and the potential use
forest,
pressures.
prefer-
address reliance on
will
cultural areas for sustainable
been minimaL Tourists
obtain correct authorization
throughout the country and hold-ups
regular military blockades
2U>
gorilla
was shown
it
may
prove tedious.
at
cies
need
to
native protein sources to
and
to
trade.
Guinea
The development
largely
due
to provide alter-
workers and
prevent their involvement
is
their families,
the
in
bushmeat
of the trade in Equatorial
to the
absence
of
sources. Research and development

fisheries,
bushmeat by
target the source of
working with logging companies
other protein
in
sustainable
domestic livestock, and other protein
alternatives might reduce demand.'
AFRICA: EQUATORIAL GUINEA
human
minimum
National awareness
All
Environmental studies and conservation education
core zones of the park while managing areas
need
around the buffer zones. '^ Training and manage-
to
be introduced
(Living Earth],
its
into the national
curriculum.
work
rangers
Mum
the country.
and carry out conservation

It
been suggested that education programs be
has
to
the government were to sign and ratify the
If
UNESCO World
proposed as a
inspire national interest
Awareness programs are also required
areas.'
growing community
expatriates,
of
the
who do
not
always realize that buying orphan apes fuels the
Logging
market
Attention
in wildlife.
cies also
of
need
conservation
International conservation agen-
made aware
be
to
of
ape
Monitoring
survival.
intensities,
At all times, the local population
integrated
management
the
in
paid to
of
ways
in
of
natural regeneration
determine sustainable
the country, thereby encouraging national interest

in
must be
Heritage Site to
and pride
the protected
making timber
extraction compatible with biodiversity conservation.
the current state
Equatorial Guinea and brought to
in
in
other protected areas around
focusing on sustainable use of forests and wildlife.

for
to a
World Heritage Convention, Monte Alen could be
intro-
radio and television stations,
to the state
must be kept
Monte Alen NP should resume
in
and be extended
both urban and rural areas.
in
of
currently seeking funds to extend
is
to Rio
education
duced
ment
nongovernmental organization, Tierra Viva
local
activities
must be
ation
of protected areas.
felling
is
needed
cycles,
to
extraction
and harvest methods, and habitat restor-
may be needed
in
buffer zones around the
parks.
FURTHER READING
Fa, J. (1992) Conservation
in
Equatorial Guinea. Oryx26
121:
87-102.
MAP DATA SOURCES

Map
16.8 Great
Butynski, T.M. (20011 Africa's great apes.
Stevens,
Arluke,
E.F..
Washington, DC.
A., eds.
In:
T.S.,
T.L.,
Norton,
B,,
Rowan,
A..
pp. 3-56.
With additional data by personal communication from Kumpel, N. 120041 and from the following sources:
Garcia, J.E., Mba,
J.
(19971 Distribution, status and conservation of primates
in
Monte Alen National Park, Equatorial
Guinea. Oryx3^ 111:67-76.

Ghiurghl, A., Puit, M. (20041 Inventaire des grands et
Monte
Alen. Rapport Technique.
Gonzalez-Kircher,
J. P.
(19941
moyens mammiferes dans
ECOFAC, AGRECO, SECA, CIRAD
('extension
sud du Pare National de
Foret.
Ecotogia y Conservacion de tos Primates de Guinea Ecuatorial. Ceiba Ediciones,
Cantabna.
Jones, C, Sabater
(Blumenbach)
Pi, J.
in
(1971) Comparative Ecology of Gorilla gorilla ISavage and
Rio Muni.
West
Africa. Bibliotheca
Wyman] and Pan
Larison, B., Smith, T.B., Girman, D., Stauffer, D., Mila, B., Drewes, R.C., Griswold, C.E., Vindum,
K.,
Nguema,
J.,
Henwood,
Research, University
L.
(1999) Biotic Surveys of Bioko
of California,
troglodytes
Primatologica 13: 1-95.

J.V.,
Ubick, D., O'Keefe,
and Rio Muni. Equatorial Guinea. Center
for Tropical
Los Angeles. Submitted to Biodiversity Support Programme. http://www.
ioe.ucla.edu/CTR/reports/CARPEl.pdt, Accessed
November 25
2004.
ACKNOWLEDGMENTS
Many thanks
Florida],
to
AUard Blom (WWF-US), Noelle Kumpel (Imperial College London], Matthew Shirley (University
John Fa IDurrell
comments on
Wildlife Conservation Trust],
Thanks
to
and Lise Albrechtsen (University
Edmund McManus (UNEP-WCMC)
for
of Oxford] for their
of
valuable
research into the literature.
AUTHOR
Brigid Barry, Tropical Biology Association
347
World Atlas
REPUBLIC OF GABON
Ambrose
Kirui,
Lera Miles, and Julian Caldecott

The Republic
Gabon
of
world's gorillas (then perhaps 35 000 animals) and
Congo Basin and the Gulf
Guinea, bordering
of
Cameroon and Equatorial Guinea to

Congo to the east and south. It has a
of
257 667
mean temperature
of
ranging from
mm
mm
3 200
estimated
area
humid equatorial with
Is
27C and an annual

the south to
in
the north.^ "
in
to
400
the north, and

total land
with an additional water area of
knn',
kml' The climate
10 000
between the
located
Is
more than
2000, forests were
In
cover 84 percent
an approximate annual loss
rainfall
country with
of the
approximately
species (Table
recorded
16.31.
at
Although both species are
throughout
the
recent years, which renders the Im-
in
populations
Map 16.9 overoptlmlstlc. Many

may have been reduced to such a low
density
in
areas affected by hunting and Ebola that
their viability
Is
threatened.
human
1.4 million (150
population
000
was
whom were
of
The major threats
ape populations
to great
mechanized logging, and the Ebola

complicating pressure of
official
language
is
French, and the
main ethnic groups are Fang, Bapounou, Nzebi, and
Up
Obamba. Gross domestic product (GDPl
that
billion,
IGNI]
2002 was
with a relatively high gross national
US$3
per person of
110, six times
higher than the sub-Saharan average.'
Abundant natural resources,
''
help to explain the country's relative
all
economic
prosperity.
played a role
in
Political
stability
has also
the country's prosperity: following
independence from France
in
1960 there have been
gorilla
human
virus, with the
population growth.
beginning of the 1980s
was thought
It
and chimpanzee populations were
relatively stable," but nest
surveys carried out be-
half,
largely
due
to
The Minkebe NP
northeast Gabon
in
data, the high frequency of
by researchers
in
encounters with gorillas
new
multiparty system and a
Introduced
elections
In
in
were
the early 1990s, and there were local
2002-2003 with a presidential
scheduled for 2005. There
because
constitution
is
some
of the long leadership tenure.
priorities include lowering
poll
political tension
Government
dependence on the
oil
industry, as the current petroleum fields are pro-
jected to run out

to
in
about 2015, and there are plans
develop other areas of natural resource ex-
ploitation
such as
1990 estimates indicated healthy populations, but
human
few
gorillas, despite the large
disturbance." The
per nesting
1998-2000, and the
number
obvious
of
mean number
declined from 6.0
site
of
1990
in
same time
one
of sites with only
the
In
period." This population collapse has
been attributed
to Ebola,
which has caused signi-
ficant mortality in both gorillas
The problem was

of
nests
to 2.0 in
nest increased from 20 percent to 60 percent
and chimpanzees.
originally restricted to the forest
Minkebe and adjacent parts
hoped that the
forestry."
and
this forest during the 1980s,
later surveys found
1967.
Illus-
trates the Impact of this viral disease. Pre-1994
area of suitable habitat and absence
power since
in
1994 and 1996.'"
Omar Bongo
in
bushmeat
hunting and Ebola hemorrhagic fever outbreaks
only two presidents, the current President El Hadj
having been
Gabon
tween 1983 and 2000 showed that ape populations
Industry accounting for 50 percent of Gabon's
GDP
until the
had declined by
a small popu-
considerable foreign support, and a booming
lation,
oil
in
In
are Illegal commercial hunting, rapid expansion of
per year' The
Income
are
may have
pression given by
expatriates], growing at a rate of about 2.5 percent
US$5
there
country,
worrying signs that ape populations

collapsed
Gabon's
All
one great ape
least
THREATS
Gabon's
2004,
national parks contain
100 km' between
of
1990 and 2000."'

In
around 64 000 chimpanzees.'""
of
Congo, and
It
was
rivers bordering the area served
as natural barriers to infected animals." The appar-
ent continued spread of Ebola
Gabon has about
however,
species, and 190
are the western

gorilla]
348
000 plant species, 446 bird
mammal
species.^
Among them
lowland gorilla [Gorilla gorilla
and central chimpanzee [Pan troglodytes
to
makes
this
In
Congo
Meanwhile, the timber industry

to the oil sector
Initially,
logging
in
2004,
view optimistic."
as an economic
Is
second only
activity In
was concentrated along
Gabon.
the coastal
Gabon was
areas, but the opening up of the TransGabonals
hold approximately 40 percent of the
railway crossing the country from east to west has
troglodytes].
thought
In
the
1980s-1990s,
AFRICA: Gabon
formerly remote parts of
facilitated accessibility to
central Gabon.'' "
The concession area
for logging
forest originally slated for logging to the core.'
increased sevenfold between 1957 (16 000 km') and
1999 (119 000 km'l. Two thirds
were logged during
methods
logging
canopy."
forest
Burseraceael
species
In
is
of
Gabon's forests
period,' using
this
selective
in
as
granted within reserves
logging
for
Is
of
found
concessions.
In
the past.' The govern-
ment now seems committed, however,
to enforcing
granted within the Lope Reserve emerged
were challenged,
particularly by the
in
1996,
European
Union program. Conservation and Rational Use

Forest
Ecosystems
which was Investing

1997, the
Central Africa
in
government
it
of
(ECOFACI,
in
conservation at Lope.
of
Gabon agreed
to
In
define a
well protected core area of the reserve, and
2000
in
allocated a less biologically rich area to a logging
company and added
of
bushmeat. An estimated
ees consumed up
In
large
200 employ-
80 tons of bushmeat per
to
Gabon.'
central
a logging
camp
these circum-
stances, hunters have a large and regular clientele,
and can make a transition from subsistence

profitable
proved
a smaller area of old growth
Table 16.3 National parks of Gabon
infrastructure
created
by
logging
the
companies has also made Gabon's urban markets
more
a
accessible. Hence,
bushmeat consumption
serious threat to ape populations
little
weight, being scarcely enforced.
although
some people
In
Moreover,
southwest
the
country are averse to eating ape meat.

sidered a delicacy elsewhere and
after, especially in
is
It
the
of
is
con-
much sought
urban areas.''
Human encroachment and
habitat
frag-
mentation has also affected the social behavior
In
White, carried out at Lope before the Increase

protection status. Indicates that logging
is
Area (km^)
Percent terrestrial
Great apes present
100,0
Gorillas
and chimpanzees
Minkebe
567
100.0
Gorillas
and chimpanzees
Ponqara
930
84.3
Akanda
538
55.7
069
100.0
Gorillas
and chimpanzees
680
100.0
Gorillas
and chimpanzees
049
100.0
Waka
Birouqou
Plateaux Bateke
Loango (comprising the former

Iguela and Petit Loango Resen/es]
Moukalaba Doudou
Presidential Reserve
and chimpanzees
Chimpanzees
Chimpanzees (orphan qoriUas)
100.0
Gorillas
and chimpanzees
196
100.0
Gorillas
and chimpanzees
165
1000
Gonllas and chimpanzees
972
6.5
Gorillas
and chimpanzees
5 672
100.0
Gorillas
and chimpanzees
152
100.0
Gorillas
and chimpanzees
Wonga Wonque
The Gamba complex
C^)
3 003
Mayumba
Gamba complex"
Gorillas
M'passa- (I'passa-I
Makokou Biosphere Reserve)
in
asso-
4 849
Mwaqne
of
Gabon. Research by
Lope
Monts de Cnstal
is
Gabon,
in
18,25,24,27,31
Park/reserve
Ivindo (inc luding the
to
commercial hunting. Second, the im-
against which the laws protecting both species have
the national parks.
Plans to log a concession that had been
but
bushmeat trade
employees consume
In
allocated
In
its
near Lope
widely exploited
is
area system, logging concessions were
logging bans
First,
year (67 kg per person per year)
Though the government has expanded the protected
Gabon.
Okoume [Aucoumea klaineana,

one of the most common tree
plywood." Most forests where the species
been
in
amount
commercial purposes, Including the production
have
The industry has created two contributory

factors for the development of the
that destroy 5-30 percent of the
Gabon and
"
Lope has since become a national park.
4 496
100.0
Gorillas
and chimpanzees
500
1000
Gorillas
and chimpanzees
is
made up
of the
Loango
NP and Moukalaba Doudou
NP, with a reserve matrix between them,
349
World Atlas
Map
in
Gabon
this cour)try profile
IWE
Gtilj
CAMEROON
of Guinea
ATLANTIC
J:
OCEAN
EQUATORIAL
GUINEA
Pd
^\\
Pongara
>
'
'
C/
Monts de 9
Crista!
NP
fvfakol<ou
GABON
,0
M'passa-
/~5
'"i---^^--'
NP
O'N
Wonga- Wongue '-^

Presidential
<
Reserve &RS;^
Lambarsne
Lac
\
v
La gun e
Nliomi Loango
ji
',;
ag u c
N do go
L
2S
Kaulomoutou
Onaiigue
Iguela
NP
HA /
Monts Doudou
^MA
<^Bimugoj
II
11
V\tfÂMBA
Petit
Loango
WcOMPLE
^ Species
Central
chimpanzee observed
Western lowland
gorilla
Estimated range
d.-j' Central
chimpanzee
Western lowland gonlla
350
after
1983
AFRICA: Gabon
ciated
with territorial conflicts
anzees
in
may
which four out
of
every
among chimpchimpanzees
five
Cama
Nature Reserve
000 km') also contains
(2
chimpanzees.
die."''
International partnerships
The international conservation and sustainable
Legislation
development community
Gabon has acceded

on
Diversity
Biological
Convention
to or ratified the
UN Framework
11997),
Gabon.
supporting a
is
number
partnership with the government
of projects in
Some
of
these are outlined here.
of
Convention on Climate Change (19971, Convention
on Wetlands
119871,
Importance IRamsar]
of International
and Convention on International Trade
Endangered Species
(CITES!
(19891,
and
1983
party to both the
is
Fauna and Flora
Wild
of
in
Timber Agreement and the
1994 International Tropical Timber Agreement.
The Gabon
demonstrates a
man-
the promotion of
management
of natural
in
ensure
resources
km'
the designation of 80 000
production forests and 100 000 km' of rural

forests,^
as well as adherence
to
of
com-
both the
management
of
and involvement
new EU-WWF network
WWF
also involved
is
in
protection and taxation
legal
management
in
for
the forestry and wildlife
sectors have been reviewed. Under
of
its
main partner, the
Water and Forests. They
moist forest as the main focus
complex
of protected
Gamba
and
agreements including the Central
in-
Africa
and
ation,
development
Declaration, and the Yaounde Declaration.^
The
applies, having recently
been replaced by
Forestry Code. Gorillas and chimpanzees are

fully
new
now
protected species under Gabonese law, having
in
the
Moukalaba Doudou NPs) has funding from US

and German agencies.'"'
982 no longer
rational exploit-
program
Protected Area Complex (Loango and
Protected Areas Network (RAPAC), the Brazzaville
transborder
Congo, and Cameroon,"
Gabon subscribes
to subregional, regional,
of a
areas between Gabon,
The integrated conservation,
of
WWF
Development (USAIDl,
has been executing the Minkebe Conservation
gazetted 5 665.5 km' of the Minkebe tropical
arrangements
The former conservation law
Gabon. Since
ment Cooperation (DGIS) and the US Agency

for International
Law 17/i6/PR/MEFCR, Gabon has set up a wildlife

management service and an antipoaching service.
ternational
of
1997, with funds from Netherlands Develop-
Project together with
Forest Partnership Convergence Plan.
extensive protection
government
efforts with the
Gabonese Ministry
Basin
of
partnerships on sustainable forestry.""
Moist Forest Ecosystems) and the Congo
and
2003
in
Minkebe and Gamba areas,

in
Brazzaville Process (Conference on Central African
Institutional
since 1992."
Ministry of Water and Forests, supporting the
to
have produced a protected area system totaling
munity
NP
Lope
of the
Global Conservation Organization
under-
determination
political
biodiversity conservation. Recent efforts to
km^ and
and development
WWF-The
has a Gabon branch, whose objectives
natural resources, and to play a leading role
iO 000
habituation
gorilla
program, and has supported the management
included policy implementation support to the
of wildlife resources,
the sustainable
several ecotourism development
including
projects,
and Forestry Policy (1992)
Wildlife
take the planning and adequate ecological
agement
ECOFAC has
long-term
research
center.
Station
d'Etudes des Gorilles et Chimpanzes (SEGCl
Lope
forest
NP
in
investigates various aspects of the
ecosystems ecology and dynamics as
well as working on gorillas and chimpanzees.
had temporary protected status since 1981 under
Global Forest Watch works with local organiza-
the previous law.'^
tions to collect
The Gabonese Ministry

is
responsible for the
resources
in
of
Water and Forests
management
of
natural
forest
The
distribute information on
been active
Gabon.
and
developments and logging impacts.'
in
(WCS) has
Gabon since 1985 and currently
runs a major country program that includes

Protected areas
activities in all the national parks, institutional
Gabon now has 13 national parks covering almost

11
at
percent of the country, and

least
support for the National Council for National
all
containing
Parks (CNPN) and the Ministry
'
The Sette
and Forests, nationwide monitoring
one great ape species."'
of
Water
of
the
351
World Atlas
bushmeat
co-funds
in
new
trade,
protected areas plan-
WCS
and reduced impact logging.
ning,
ttie
SEGC long-term
used by both
in
fessional
national parks. Other priorities include legislative
Lope,
and the
forestry school
ttie
Group. The overall priority
researcln center
Lope and runs a training center
enforcement and the development
national university."
The
in
actions to halt the decline of great apes
km' Mpassa Biosphere
the 100
in
its
1961 and
in
city to
has been recently refurbished
trade,
the oldest field research station
the country.
It
EU and
with funds from the
equipped
is
regional and
implications, increased understanding
manage
protected areas and the
of wildlife
departments needs
be explored, and laws governing the conduct of
national
logging companies need to be applied.
workshop was organized

Sanctuaries
Projet
be
to
strengthened, alternative sources of protein need to
conservation network.
The
and capa-
bushmeat
and further species population surveys.
The capacity
newly created Ivindo National Park, are an

part of the
Gabon
Bushmeat trade and use
and Mpassa Reserve, now within the
integral
in
to
The research
receive students and scientists.

station
national
include the promotion of awareness on hunting and
Reserve near Makokou, was established

is
of
strategy for the use of wildlife resources. Priority
de Recherche en Ecologie Tropl-
Instltut
cale (IRETI,
Programme and Primate Specialist

now is to develop promanagement capacity in all of these
Tropical Forest
also
A bushmeat
2002 by the Ministry
in
of
Water and Forests, and placed an emphasis on

des Gorilles (PPG)
Protection
Plateaux Bateke
NP
Aspinalt Foundation
orphan gorillas
is
the
education, training, and awareness-raising, both for
supported by the John
the general public via information campaigns, and
(JAFI,
the
into
and aims
wild
Sanctuary on the Bateke Plateau.'
to
for
working
staff
evaluation,
management."
wildlife
in
recommendations
specific
currently
is
It
release
Mpassa
the
at
in
the creation
of
included
an
office
Its
resource
for wildlife
holding about 20 gorillas for this purpose. The Petit
management, employment
Evengue program run by Operation Loango has
creation of a joint action plan, and identification of
gorillas. ''
The Societe d'Exploitation du Pare de
six
provincial
of
guards,
funding sources.
la
Lekedi (SODEPALl has also provided sanctuary for
chimpanzees and
Ebola virus
gorillas.'"
Since the Ebola virus poses such a threat to apes
Gabon, an effective plan
Overview
priority.
The creation
line
with
of 13 national
the
parks
for
priorities
2002 was
in
Gabon
of
combat
it
is
in
an urgent
plan would need to address in-
tensified research on reservoirs
in
lUCN-The
World Conservation Union, as recommended by
Such
to
and hosts
of the
disease, including their ecologies, as well as on
vaccines and their possible
its
modes
of deployment.^"
FURTHER READING
Collomb,
J.G., Mikissa, J.B.,
Akagah,
S.,
Resources
Fragmented
Gabon. Primates UQ
Fernandez,
Newsletter
3:
E.,
S.,
Mofouma,
Mundunga,
A. (20001
S.,
Nzao Nzao,
First
H.,
Mapaga,
Look at Logging
in
J.,
Mikolo, C, Rabenkogo, N.,
Gabon. Global Forest Watch, World
Washington, DC. http://www.globalforestwatch.org/common/gabcn/english/report.pdf.
Institute,
Tutin, C.E.G. (1999)
Tutin, C.E.G.,
Minnemeyer,
Bayani-Ngoye,
[II:
1^.
living:
behavioural ecology of primates
in a
forest
fragment
in
the Lope Reserve,
2i9-265.
(1983) Gorilla and
Chimpanzee census
in
Gabon. lUCN/SSC Primate Specialist Group
22-23.
MAP DATA SOURCES

Map
16.9 Great
Butynski,
TM.
(2001) Africa's great apes.
Stevens, E.F, Arluke,
A., eds.
In:
Beck, B.B., Stoinski, tS., Hutchins, M., Maple, TL., Norton.
B.,
With additional data by personal communication from Courage,
352
Rowan,
A.,
A. 12004)
AFRICA: Ghana
Barnes, R.F.W., Jansen, K.L 119871 Forest Elephant Survey, Progress Report
Conservation International INYZSI. Conservation International.
Blom,
Barnes, R.F.W., Barnes, K.L. 119921 Primates
A., Aler, P.T., Feistner, A.T.C.,
Oryx 26
distribution.
Gabon: current status and
in
223-23/.,
141:
lUCN Conservation Monitoring Centre
UK and
1986-1987. Report for Wildlife
New York.
lUCN Directory
(1985) Ttie
of Afrotropical Protected Areas. lUCN,
Cambridge,
Gland, Switzerland,
Fernandez, M. (1984) Nation-wide census
Tutin, C.E.G.,
troglodytes] populations
in
of Gorilla
[Gorilla g.
Gabon, American Journal of Primatology
White, R.J.T. 119941 Biomass of rain forest
mammals
in
6:
gorilla]
and Chimpanzee [Pan
t.
313-336,
the Lope Reserve, Gabon. Journal of Animal Ecology^!. 345-367.
ACKNOWLEDGMENTS
Many
thanl<s to
Lee White
John Ady lUNEP-WCMCI, Allard Blom IWWF-US], Sally Lahm (University

Conservation Society! for their valuable
(Wildlife
comments on
of California,
San
and
Diegol,
AUTHORS
Caldecott. UNEP World Conservation Monitoring Centre
Ambrose
Kirui,
Lera Miles,
Julian
REPUBLIC OF GHANA
Ediviund

The Republic
West African
It
Ghana
Ghana's
situated on the southern
is
coastline, bordered by
Burkina Faso
west.
of
to the north,
has a land area
composed mostly
of
Togo
and Cote
of
McManus
to
the east,
d'lvoire to the
239 460
km^ and
low plains, with a plateau
south-central area. The climate
is
generally
in
is
the
warm,
developed close
Lieutenant Jerry Rawlings
Council,
way
savanna bush
to
more northern Brong-Ahafo
region.
In
in
the
the south-
western areas where most chimpanzees are found
(Map
16.101,
two rainy seasons occur, from May
July and from
September
Ghana has
estimated
in
2003
was once home
a
to
diverse
powerful AshantI kingdom,

of the British colony of
the Gold Coast through a series of wars

century. In 1901, the Ashanti
Territories Protectorate
the 19th
kingdom and Northern

into the
German Togoland was
placed under Gold Coast administration
state to achieve full
in
were amalgamated
Gold Coast, and neighboring
957, the Gold Coast
population,
be about 20.3 million.'" The area
to the
became Ghana,
the
in
the
1980s.
early
of the ruling Provisional National
abolished
the
Defense
dissolved
constitution,
parliament, and outlawed the opposition political

parties.
The ban on opposition parties was
the mid-1980s, and a
new
constitution
up and approved by referendum
two presidential elections,
in
in
lifted
by
was drawn
1992.
The
first
1992 and 1996, were
endorsed by international observers, and returned
to October.'^
culturally
which resisted the expansion
to
first of
Rawlings seized power by force, assumed chair-
manship
This dry forest gives
1966, the
in
several that ended with the ascendancy of Flight
many moist forest fragments,
which become progressively drier towards the north.
He was
ties with the Soviet bloc.'
overthrown by a military coup
with a strong moisture gradient. The southwestern

part of the country has
Kwame Nkrumah, was
ruler,
first
the pioneer of 'African socialism' and gradually
in
first
1919.
In
African
independence from the UK.
President Rawlings to power He stood

after a constitutionally limited
and was replaced through the election

leader John Kufuor.
In
down
two terms
December
in
2000
of office,
of opposition
2004,
Kufuorwas
re-elected for a second four year term of office.
The international community perceives Ghana

as very
much
pluralistic political
of power.'
'
inherited an
measures
success
story,
in
terms
of
its
system and smooth transitions
The Kufuor administration, however,
economy
in
decline,
and took strong
that included the raising of fuel duties by
353
World Atlas
Map
16.10 Chimpanzee distribution
in
Ghana
2W
BURKINA FASO
this
country profile
0?E
^^^^T'C:^^-
Species

Westem chimpanzee observed
after
1983
Western chimpanzee
'"4]
Western chimpanzee presence alleged
=?
^^
BENIN
Bunkpurugu^^^
1940
Estimated range
C^
Western chimpanzee
Tamale
(N^
^_.
c6te
DM VOIRE
TOGO
.,-'
GHANA
BRO N G - /vQa^FO
F
Lake
Volta
Bia
/ Ho
pKoforidua
Yoyo River FR
Tano-Nimiri
.,
FR
Boi-Tano FR
J
Accra
'Nini-Suhien
NP
Ankasa
-AKOSOMBO
DAM
FR
Dadieso
Cape Coast
River
A TL
ANTIC
OCEAN
RR
Draw
FR
Cape Three
354
>
MP & BR
Points
FR
AFRICA: Ghana
over 90 percent.'"
In
2001, a recovery
gold and
in
cocoa prices helped Ghana attain macroeconomic

stability,
and
'heavily indebted
poor country' (HIPCI status by the
making
International Monetary Fund,
forms
certain
declining, but
was granted
2002, the country
in
debt
of
eligible for
it
poverty
Overall,
relief.
is
locally extinct in
Kakum and
Cape Three Points." No chimpanzees were detected
the Yoyo River or Dadieso Forest Reserves,
in
although anecdotal information from hunters suggests that they are present."
unemployment remains high and food
crop farmers remain vulnerable.'" The govern-
THREATS
ment's economic and social policies are consistent
All things
w/ith
may be
Reserves." Chimpanzees
several other forest areas including
international development targets, but
Ghana
considered,
is
it
anzees are nearly extinct
possible that chimp-
in this
country. Hunting
is
" and
it
remains dependent on international financial and
a significant threat to wildlife in Ghana,'"
technical assistance.
is
About 57 percent
of the country's land
area
devoted to agriculture, particularly cocoa and
palm
plantations.'
is
areas, Nini-Suhien NP, Bia NP, and Boi-Tano and
By 200^, subsistence agriculture
Dadieso Forest Reserves have been subject to

particularly heavy hunting pressure.' " Primates
gross domestic product
of
(GDPl and employed 60 percent of the work force,
Some
mainly small landholders.'
300 000 people
work as hunters.'^' There
is
harvest of 385 000 tons of
bushmeat
an estimated annual
118.8
kg per
person per year) worth about US$350 million,
which around 60 percent
sold
is
in
of
urban areas."'
and cocoa production are major
timber,
sources
of foreign
bushmeat when they can." Of the protected
eat
oil
accounted for 35 percent
Gold,
estimated that 90 percent of Ghana's population
make up
animals
a
major factor behind the extinction
watdroni].'"
anzees
land
in
chimpanzees [Pan trogiodytes

limited
to
southwest
the
of
they are probably found only
They are
verus].
the
in a
country,
where
few moist semi-
The pet trade
Ghana."'
In
addition,
in
lost
much
of its forested
200 km'
(1.7 percent).'
The
major pressures on forest have been road construction, agriculture,
and timber extraction, with
mineral exploitation an emerging threat around the
They may also occur
to
dry semideciduous forests, but this has not been
also thought to have
remain by 2000, with an
to
average annual loss of
year 2000." "'"
some moist evergreen and
is
the 2Gth century.' Around 63 350 km' of
deciduous and wet evergreen forest fragments.

in
Miss
^'
Ghana
was thought
forest
Ghana are western
of the
affected the remaining wild populations of chimp-
In
the
of
and traded: hunting pressure was
Waldron's red colobus monkey [Procolobus badius
exchange.
The only wild great apes found
small but significant fraction
killed
Demand
major degradation
for land
and timber has
led
and encroachments on
of
protected areas."
confirmed."
The most recent population estimate was
made
in
when
1995,
500-2 200 chimpanzees were
thought to be present
were thought
to
in
Ghana."
In
1979, there
be fewer than 200;" by 1988. the
estimate had risen to 300-500 chimpanzees.^' These

increases are a result of improved knowledge; the
actual
number
chimpanzees has almost
of
certainly
Extensive surveys were carried out

'
Chimpanzees may
still
in
1999 and
be present
in
the
78 km^ Bia National Park INP] and the 160 km'
Nini-Suhien NP, but none were detected by these
surveys
in
either location.
National laws and protected areas
Chimpanzees were wholly protected under the Wild

Animals Preservation Act
Wildlife
Chimpanzee presence
was confirmed in the Ankasa Resource Reserve

1343 km'l," Draw River Forest Reserve 1235 km'),
(Act
43
of 1961),
strengthened the legal protection
The
Wildlife
and Forest
of
of
chimpanzees.
Policy, introduced in 1994,
promoting sustainable forest
management.'^ Penalties
ging,
and the
Conservation Regulations 11971) further
had the aim
fallen over the period.
2001.^'
for illegal hunting, log-
and other forest crimes were increased
in
2002."
Bia
NP was
was reduced
to
gazetted
78 km'
in
in
1974
at
306 km', but
1976, with the remainder
and the Tano-Nimiri Forest Reserve 1205 km'].
of the park becoming a resource reserve with

timber concessions.'' " Bia was designated as a
Biosphere Reserve in 1983." " Nini-Suhien was
Chimpanzees are
designated as a national park
Hills
1295 km'l
likely to
occur
in
the Krokosua
and Boi-Tano 1128 km'l Forest
Most
of
in
1976."
the remaining closed forest
is
found
in
355
World Atlas
the 18 000
km'
production
reserves where timber harvesting
km'
permitted, and U 500
where
tended
ance
Is not.'''
It
of
Ankasa Conservation Area lAnkasa Resource

Reserve combined with the adjacent Nini-
Is
Suhlen NP) and the Krokosua
reserves
protective
" The protective reserves are
Reserve. Additional surveys
In-
safeguard water bodies, areas of import-
to
and
for biodiversity,
addition, the
new
much
baseline of chimpanzee status
biodiversity conservation
in
support of
and alternative
livelihoods.
develop a
IGEFI funds
in
exchange
recorded
scheme covers Dadieso
other
conceded
for logging in 20011,'*
of
Draw
in
initiated in 1994, but
have been
killed by
1972.^
chimpan-
to
low
human
more frequent than
capacity development
Afurtherattempt was
populations
as
hunters."
In
the form of training of
staff. In
order
to
of local
fication of
The West African Chimpanzees: Status Survey and
made
for priorities in
the
Reserve
following
staff
Ghana."
the
chimpanzee range
many
in
field
identi-
measures. As
West Africa
is
and
coordinated national cooperation and com-
mitments
reserves, to improve forest
to
chimpanzee conservation
will
be
needed. The Ghanaian government has stated
to collate basic ecological
that protocols for cooperating
Research and surveys. Estimates are needed
ment
chimpanzee numbers and
in
countries, joint
data.
of
pro-
should complete their training.
Transfrontier conservation
Overall priorities. Major efforts are needed to
management, and
in
chimpanzees, and Ankasa Resource
continuous over
protect existing
hunters
improve monitoring
training Wildlife Division staff
recommendations
In
tected areas there should also be a focus on

Plan
in
and
in
any other area
in
park rangers and recruitment
these animals are believed to
Conservation Action
likely site to
Ghana. This area should receive Immediate
rehabilitation attempt concerning six
Bia NPfailed
most
around the reserve), with chimpanzee sightings
in
plans for chimpanzees and
identified as the
Ghana Idue
zees
system
augment the development
to
support a sizeable chimpanzee population
River and Krokosua Hills Forest Reserves.
a geographic Information
wildlife.
has been
was mostly
and parts
to
Protection. The Krokosua Hills Forest Reserve
Forest
Reserve, which has never been logged," part of Yoyo

River Forest Reserve Ithe remainder
In
management
of
these areas." Of those areas relevant to great ape

conservation, the
be used
year conservation program. All
database
IGISI
logging
for not
five
sufficient
Ghana. The
chimpanzee habitats should be mapped and
Local communities will receive Global Environment

Facility
In
results of these surveys could
of the forest reserve area, mobilizing
national and international resources
Yoyo River
Reserve would probably provide a
intended to prevent logging
Is
Forest
Hills
In
Forest Reserve, Bla NP, and Dadieso Forest
fragile ecosystems.'^ In
globally significant biodiversity
area IGSBA] system

over
which Include
of forest reserves,'
of
transboundary
In
the develop-
protected
areas,
including for Bia NP, are to be discussed."
distribution In the
FURTHER READING
Grubb,
Jones,
P.,
T.,
Davles, A., Edberg,
E.,
Starin,
E.,
Hill, J.
(19981
lâmmals
of Gfiana, Sierra
Leone and The
Ram
Forest: Forest
Gambia. Trendhne Press, Zennor, UK.

Swaine, M.D. (19811 Distribution and Ecology of Vascular Plants
Hall, J.B.,
Vegetation
Magnuson,
in
Adu Nsiah,
L.,
in
a Tropical
Ghana- Dr W. Junk, The Hague.

M., Kpelle, D. 12003)
Ghana.
In:
Kormos,
R.,
Boesch, C, Bakarr,
M.I.,
ButynskI, T.M., eds.
West African Chimpanzees: Status Survey and Conservation Action Plan. lUCN/SSC Primate Specialist Group.
lUCN, Gland, Switzerland, pp. 111-116.
Gates,
J.F.,
Abedi-Lartey, M., McGraw, W.S., Struhsaker,
colobus
monkey Conservation BiologyM
(51:
T.T.,
Whitesides, G.H. (2000) Extinction
of a
West African red
1526-1532.
MAP DATA SOURCES

Map
356
16.10
following sources:
AFRICA: Guinea
Butynski, T.M. 120031 The chimpanzee Pan troglodytes: taxonomy, distribution, abundance, and conservation
status.
In:
Kormos,
Boesch, C, Bal<arr.
R.,
M.I., Butynsl<i, T.M..
West African Chimpanzees: Status
eds,
Survey and Conservation Action Plan. lUCN/SSC Primate Specialist Group, lUCN. Gland, Switzerland.
pp. 5-12.
Magnuson,
Adu-Nsiati, M., Kpelle, D. 120031 Ghana.
L.,
In:
Kormos.
Boesch, C, Bakarr, ML, Butynski, T.M., eds.
R..
West African Chimpanzees: Status Survey and Conservation Action Plan. lUCN/SSC Primate Specialist Group.
lUCN, Gland, Switzerland, pp.
1 1 1
1 1
ACKNOWLEDGMENTS
This country study draws extensively on the
Ghana chapter from the lUCN/SSC West African Chimpanzees: Status
Sun/ey and Conservation Action Plan.^' Many thanks
comments on
to
Lindsay Magnuson [College of the Redwoods! for her valuable
COMPILER
Monitoring Centre
REPUBLIC OF GUINEA
Muhammad Akhlas

The Republic
of
Guinea
and 2003. Unrest
located
is
on the West
African coast, with Sierra Leone and Liberia to
spilled over into
in
Sierra Leone and Liberia has
Guinea on several occasions over
tfie
the past decade, threatening stability and creating
south. Cote d'lvoire and Mali to the east, and Senegal
humanitarian emergencies. This has undermined
and Guinea-Bissau
investor confidence and the ability of the country to
the north, and has a land
to
area of 245 857 km^.
Its
clinnate is hot
and humid
but seasonal, with a rainy season from June to
Novennber and
country
is
a dry
season
at
other times,
the
affected by dry, dusty harmattan winds off
the Sahara. Guinea's terrain comprises a
an undulating
plain with
In
when
coastal
flat
mountainous
to
out of poverty The International Monetary
Fund IIMFl and World Bank cut

in
strengthen
to
off
but economic activity
2003,
in
most assistance
was expected
to
2004 as the security situation started
improve.^
interior.
was estimated
2000, total forest cover
move
to
be
69 290 km^ or 28 percent of land area,' with about
Guinea has about 15 species
17 percent 111 821 km'l of
the western chimpanzee [Pan troglodytes verus]
it
within 162 classified
of primates, of
There appears
which
forests. In 2003, 3.6 percent of the terrestrial area
is
was
reasonably healthy population that amounts

36-51 percent of all western chimpanzees." "
cultivated
crops.^
and 2.4 percent under permanent
The remaining area
The population
million
people
in
of
Is
woodland savanna.
Guinea was about 9.25

growing
200'!,
on a mixture
people that
of agriculture,
live in
bauxite
(aluminum
ore]
rate of 2.4
is
based
which supports those
rural areas,
has more than 30 percent
at a
economy
percent annually^ The country's
of
and mining. Guinea

the world's
accounts for about 75 percent
of its exports by value.
The country achieved independence from

France
in
a military
of that
1958, and
was
government
ruled without elections by
until 1993,
when
the head
government. General Lansana Conte, was
elected president, with renewed
mandates
in
1998
ape.
be a
to
has therefore been suggested that the country
one day be the
last
stronghold
of this
to
It
may
subspecies,"
but this will depend on the fate of the country's

forests,
in
and deforestation
is
an ongoing pressure
Guinea.
known
and the mining sector
only great
the
Chimpanzees are known

Guinea.
to
recorded their occurrence
in
be widespread
the mid-1980s
27-30
of the country's
34 prefectures.'" A study published

firmed their presence
more
at 71
in
in
questionnaire survey
sites,
in
1998 con-
and included a
detailed questionnaire survey that reported
chimpanzees
estimated
at
606 locations." This led
national
population
of
to
an
113-29 011
357
World Atlas
Map
16.11
Chimpanzee
distribution in Guinea
eo
'
this
country profile
CO
(01
IS
-t
"8
li-^
t
a
JIT"^
<
v-=.
Ci
.'^
Hi
S -So
b
^^
EM
TO
CD ofl
"/^,,.
CD
UI
en
r**
0)
o o
C3.
oj o
ci^
BS
CO
{/)
fig '^*ii
Z5
"o
i^:5^_/?
s.
Ed
7s -e
,ow 1
z
o
u
Ji
-i
+sV--
C^
<
of
m
Ed
1/1
Oo
^.
'-"'"^^:;s
r>
in
i={
Si
E^
<u
.fa^
er>
n n
-n
Cl)
^
I
n
o
n
o
ai
n
o
n
o
Q-
fl)
(1)
fl>
tn
en
<
tn
n
b
lid
Z
<
oj
m
o
b
n
b
fi
b
o
'?
F
Dl
* 5 < ^ ^ ^
358
m
n
ra
m
F
tu
AFRICA: Guinea
of
Central African Region (1981). Guinea has signed
420-6 625 individuals that arose from the 1980s
but not yet ratified the African Convention on the
chimpanzees;
1
than
greater
estimate
the
survey. Eartier estimates had suggested that there
were fewer than 12 500" and
000-10
000.^'
were not generated using comparable
figures
techniques, and so
it
not clear whether the
is
is
Conservation
Nature and Natural Resources.'
of
Chimpanzees are
These
'integrally protected'
under
Guinea's law governing the use of wildlife," and
may
not be hunted, captured, detained, or exported
without a scientific permit obtained from the
stable.
Small populations are known
to
occur
in
forest
government. Legislation
areas throughout Guinea. Important sites include
however, unless
Mount Nimba Biosphere Reseri/e
culture,
Bossou
(including
is
it
and only
in
has
typically
impact,
little
consistent with the prevailing
some areas
of Guinea,
such as
du Ziama Biosphere Reserve, Haut
the Fouta Djallon area, are local people traditionally
Niger National Park INPI, and the classified forests
averse to the hunting of chimpanzees."' " These
forest], lulassif
of
Nialama, Sala, Fello Digue, Balayan-
Diecke,
Souroumba, Bakoun, and Souti Yanfou." Population

density can vary greatly at each
NP
Haut Niger
in
in
2001,
where
gallery forests to 0.1/km^
in
it
site, for
example
ranged from
at
particular areas
are
3. 5/km'
wooded savannas.'
Protected areas
THREATS
ing
being driven by population growth
is
human and
chimpanzee populations."
There are three protected areas
Deforestation
densely peopled,
relatively
enabling a positive relationship between
chimpanzees.
strict
in
Guinea contain-
the southeast, there are two
In
nature reserves: Mount
Nimba
(130
km'),
and agricultural expansion, aggravated by the use
designated as a Biosphere Reserve and a World
of fire to clear land in the dry season.^' Selective
Heritage Site
logging
is
also a significant threat to chimpanzees,
with a range of direct and indirect impacts on forest

structure,
connectivity,
and species composition,
and on chimpanzee ranging behavior, with logged
more
forest being
likely to
be cleared by settlers."
"
(1
Ziama
1981, and the Massif du
in
123km'l, a Biosphere Reserve since 1980.
central Guinea there
Park (NPl
2002.
(6
in
the Haut Niger National
is
000 km'), a Biosphere Reserve since
The core zone
makes up about
Haut Niger NP, Mafou
of
forest,
10 percent, with the remainder
buffer zones." The
NP has
Other pressures on chimpanzee habitats include
being
mining and infrastructure development. The north-
received
ern part of Mount
support through the European Union (EU), but the
Nimba Biosphere Reserve has
significant
been disturbed by iron-ore mining,' and the expan-
other
sion of railway and road projects through protected
management and
areas
protected
Haut Niger
and management
funding
from
lack
of
Guinea are protected
for
suffer
resources.
areas has posed serious consen/ation problems.'^
Hunting also affects Guinea's chimpanzee
Commercial hunting
population.
spread,'^ though not universal."
for
meat
Farmers
is
kill
wide-
chimp-
Sacred sites
A number
anzees as crop pests, and they are sometimes
Bossou. This
hunted for the medicinal or magical properties
hills,
their
body parts
(for
example, the blood
cure epilepsy, and the meat
young
past,
children).'
is
is
of
thought to
believed to strengthen
Orphans are sold as
pets. In the
many chimpanzees were captured and shipped
overseas
for
biomedical research."
areas
of
in
known
reasons; the best
religious
site consists of
these
of
is
several small sacred
situated within a farming region of smalt
vil-
lages and fields. Yukimaru Sugiyama and col-
leagues
at the
University
socioecology,
at
Primate Research
life
Institute of Kyoto
been researching chimpanzee
have
history,
demography, and
tool
use
Bossou since 1976." The chimpanzees here are
not completely isolated from others, for occasional
visitors
Legislation
have vanished. Bossou
Guinea
which there
Trade
is
in
party to the Convention on International
Endangered Species
of
Wild Fauna and
Flora (CITESI. Guinea has also ratified the World
Heritage Convention and the Convention for Cooperation
in
the Protection and Development of the
Marine and Coastal Environment
of the
West and
have been seen and maturing individuals
transfer
is
km
one
of the
among chimpanzees."
neighboring population
6-10
is
few
sites at
evidence of male intercommunity
is in
the
" The nearest
Nimba Mountains,
away, and there are efforts
forest corridor to
link
to
develop a
these areas (the Green
Corridor Project].'
359
World Atlas
Education and sanctuaries
integrated
The Chimpanzee Protection Project IPCC) was
components
funded bytheEU
program.'
PCC
Carter. Major
ment
in
1995-1999 and directed by Janis

objectives included the involve-
chimp-
of local people, including hunters, in
forests
Balayan-
cational activities are ongoing
by
International
to
continue certain
funds from
the
activities,
US Agency
for
Development lUSAIDI and Friends
of
The educational
the
PCC and onwards
conducted during
activities
included slide programs and
bumper
the production of posters, brochures, and

stickers addressing the
tect
need
conserve and pro-
to
chimpanzees, the current legislation protecting
them, and the impact

infant.
In
mental education
these areas.'
in
Nimba
the Bossou and
of
purchasing an orphaned
Radio programs were also produced
University Primate Research institute have been
raising
Environmental
running
in
Nimba
Biosphere
nine villages
Reserve,
Appel de Detresse (Distress
Call)
entitled
was produced and
distributed throughout Guinea, particularly
prefecture of Pita
where
widely
These
supported by
In
prefectures were
visited,
distributed,
the
educational materials
meetings with elders held, and slide
programs shown.
being
in
In
2004, Appet de Detresse
and translated
revised
into
various
was
local
languages under a US Fish and Wildlife Sen/ices
lUSFWS)
pamphlets,
while
Fouta
been
have
activities
USFWS
for
to
start
of
education and
chimpanzee conservation.
activities
early 2005, and to
in
Chimpanzee
Wild
the
Djallon,
raising
financially
since 2003.'
performances and distribution
were scheduled
include dramatic
newsletters.
of
Orphan chimpanzees are received
schools and sous-
all
been
Mount
badges, and videos have been distributed more
in
capacity
schools.
have
the area of the
in
awareness
in
A brochure
16
to
sessions
education
governmental organization involved

raising.
at
awareness about chimpanzees and the
Foundation (WCFl has a program
awareness
environ-
ongoing since 1993. Since 2003, books aimed
collaboration with Guinee Ecologie, a national non-
building and
region,
coordinated by the Kyoto
activities
environment have been donated
the Animals.
for care
and
rehabilitation by the Centre de Conservation pour
Chimpanzes (CCCJ, located near the
village
of
Somoria within the Haul Niger NP, and supported
November
by the US-based Project Primate. By

2004, the sanctuary had taken
in
37 chimpanzees."
grant.
USAID
and
Winrock
International
Tatyana
Mount Nimba.
co-managed
the five
Souroumba, and Souti Yanfou, and similar edu-
supported
to
in
Nialama, Sincery Oursa,
environmental education, focusing on the areas
education and long-term monitoring
Bossou
Bakoun,
anzee population monitoring and village-scale
Subsequently, Carter was able
Green Corridor' linking
'*
'
PCC education and monitoring

their forest co-management
Chimpanzees are now monitored
into
by local residents
of
around Bakoun and Nialama Classified Forests.
Groundwork for the
the
later
Humle

In
October 2004, a National Great Apes Survival Plan
INGASPI Workshop was held
The NGASP
is
in
the capital, Conakry.
based on a previous workshop held by
Conservation International
September 2002 and
in
on the subsequent action plan
lUCN-The World
of
Conservation Union for western chimpanzees.'' This

indicates the following priorities for conservation
in
Guinea.
Research and surveys. There
is
need
for
comprehensive information on the present

status,
number,
distribution,
conservation measures
nature of and
petition
in
potential
threats,
and
Guinea, and on the

for
resolving
com-
between humans and chimpanzees
over natural resources. Studies are needed to

clarify
aspects of chimpanzee behavior and
social ecology,
and their populations
habitats; to identify practical
360
ways
to
in
various
reconnect
AFRICA: Guinea
fragmented habitats through natural forest

corridors;
and
to identify priorities tor
doing so.
Protected areas. Priority areas include Fouta
Nimba Mountains, and Haut Niger
Djallon,
Connprehensive steps are needed

standards
the
management
protected
existing
of
improve
to
planning,
protection,
of
and
areas,
on the support of the international
calling
community and the involvement
ment and
local
govern-
of
communities.
Peace and security. International organi-
may need
zations
to
be involved
Chimpanzees
to help resolve
border issues and increase security
.-
*'
trans-
in
"%
boundary protected areas.

Capacity building. There
then the capacity
management
need
is
sessment
implement national and
be paid to the distribution
and awareness-raising programs regarding
sure that chimpanzee consen/ation and local
chimpanzee hunting, bushmeat, and the pet
among
schoolchildren
include
Chimpanzee and
may
be possible
and
the
wildlife-oriented
in
some
tation
and educational use
tourism
Development. Environmental impact assess-
ments are needed
prior to the initiation of
mining ortimber extractive
should only be promoted
anzee
study
nuts.
orphaned and
confiscated chimpanzees.
be carefully regulated. Chimpanzee tourism

the
palm
for the rehabiliof
military.
areas, but should
if
oil
revenue, to en-
of
secure financing are needed
Guinea
in
hammerandanvilto
crack open
populations benefit. Finally, sanctuaries with
communities.
Suggested educational targets
Bossou
stones as
must
risks of disease transmission. Attention
needed
at
that they will not suffer negative
Education and tourism. Further education
are
of
impacts from habituation or undergo serious
international legislation.
trade
use a pair
Matsuzawa Tetsuro
to streng-
protected area and wildlife
of
staff to
.,
of
individual populations leads to a positive as-
habitat,
activities in
and guidelines
to
new
chimp-
minimize
impacts are required.
FURTHER READING
Barnett, A.A.. Prangley, M.L. 119971
preliminary check-list and a
Mammalogy
in
summary
research recommendations for the future.
of
the Republic of Guinea: an overMiew of research from 1946-1996, a
Mammal
Review Tl
13):
115-164.
Humle,
Matsuzawa,
T.,
T.
12001)
Behavioural diversity
neighbouring areas, Guinea and Cole
Kormos,
Boesch, C, Bakarr,
R.,
M.I.,
d'lvoire.
among
West Afnca:
the wild
a preliminary report. Folia
Bossou and
of
Pnmatologica 72
121;
57-68.
Butynski, T.M., eds (2003) West African Chimpanzees: Status Survey and
Sugiyama,
of
Y.,
Soumah,
A. (19881 Preliminary survey of the distribution
and population
of
chimpanzees
in
the Republic
Guinea. Primates 29: 569-574.
Yamakoshi,
G..
Takemoto,
chimpanzees
at
H.,
Matsuzawa,
T.,
Sugiyama,
Y.
Bossou, Guinea. Primate Research 15
Ziegler, S., Nikolaus, G., Hutterer, R. (20021 High
Niger, Republic of Guinea.
Oryx3i
11):
mammalian
(19991 Research history and conservation status of
(2):
101-114.
diversity
in
the newly established National Park of Upper
73-80.
MAP DATA SOURCES

Map
16.1
Chimpanzee data are based on the following sources:
Butynski, T.M. 12001 Africa's great apes.

1
Stevens,
E.F.,
Arluke,
A.,
In:
eds. Great Apes
T.S.,
and Humans: The Ethics
T.L.,
Norton,
B.,
Rowan,
A.,
of Coexistence. Srr\iH\son\an Institution Press,
361
World Atlas
Butynski, T.M. 120031 The chimpanzee Pan troglodytes: taxonomy, distribution, abundance, and conservation status.
Kormos,
Boesch, C. Bakarr,
R.,
Butynski, T.M., eds.
M.I.,
In:
Kormos,
R.,
Humle,
C, Tounkara,
T.,
Bruglere,
D.,
M-C, Matsuzawa, T., Suglyama, Y
Fleury-Bruglere.
The Republic
E.O. 12003)
of
Guinea.
In:
Kormos,
Boesch,
R.,
West African Chimpanzees: Status Survey and Conservation Action
Plan.
C.,
Carter,
Bakarr,
J.,
M.I.,
DIalLo. M.S.,
Sagno,
lUCN/SSC Primate
Specialist Group.
lUCN, Gland, Switzerland, pp. 63-76.

ACKNOWLEDGMENTS
This country profile draws extensively on the Guinea chapter from the
Survey and Conservation Action
P(an.''
Many thanks
to four
lUCN/SSC West African Chimpanzees: Status
anonymous reviewers
for their valuable
comments on
the
draft of this section.
COMPILER
Muhammad
Akhlas, University of East Anglia
REPUBLIC OF GUINEA-BISSAU
CiûDiA SouSA, Spartaco Gippoliti, and

The Republic
Mozambique, and East Timor] the country suffered
Guinea-Bissau
of
one
is
the
of
snnaUest countries on the Atlantic coast of West

Africa,
sandwiched between Senegal
and Guinea
36 125
km^
to the
it
to the north
south and east. With an area of
number
includes a
of snnall offshore
terribly,
and
is
now deeply Impoverished. depends

Cashew nut proIt
mainly on fishing and farming.

duction
is
increasing, and
comes from
amounts
with relatively small
kernels, and timber. Rice
ated from the mainland by wide intertldal mudflats.
staple food.
Guinea-Bissau's population, which includes about
much-needed revenue
was approximately 2A
million
in
200i, and
was growing
at
After Independence
from Portugal
197i,
In
Guinea-Bissau established a one-party system and a
economy A
centrally planned
established a
stance, which
elections
in
military
coup
in
1980
new system with a more pro-market

won a mandate in the country's first
1994, but there
were repeated coup
attempts through the 1980s and 1990s, one of which

led to civil
war
in
998. Intermittent fighting
Senegalese-backed
military junta
government
destroyed
much
troops
of
economy. A
brief
return
to
2000-2002 ended with another coup

2003 that
that
installed the current
and
the country's
damage
democracy
infrastructure and caused considerable
the
between
in
Offshore
in
the
long
run,
but are
Sixty percent 121
870 km^l
of
Guinea-Bissau
was forested in 2000, mainly with natural broadleaf

humid or semidry forests. The country has the
largest
Africa;
area of mangroves and coastal flats

originally
1 1
percent of the country
in
was
covered with mangroves.'

Guinea-Bissau has
11
species of wild primates, of
which the western chimpanzee [Pan troglodytes

verus] is the only great ape.^ " Due to the very
limited
survey data,
in
estimates range from 600

Until 1989,
in
uncertain
is
it
chimpanzees are found
September
peanuts, palm
of
the major crop and
reserves could provide
oil
to
government.
is
currently unexploited.^
about 2 percent
annually.^
most foreign exchange
the export of fish and seafood along
islands - the BIjagos archipelago - that are separ-
20 ethnolinguistic groups,
in
to
but
000 individuals.''
chimpanzees were thought
the country," but they
how many
Guinea-Bissau,
to
were confirmed
be extinct
to
be pre-
Like the other former Portuguese colonies
sent after a comprehensive wildlife inventory
were abandoned
undertaken by Guinea-Bissau's Direcao General das
much
362
Muhammad Akhu
preparation
in
to
independence without
the
mid-1970s
(Angola,
was
Florestas e Caca (DGFC) and the Canadian Centre
AFRICA: Guinea-Bissau
Map
16.12 Chimpanzee distribution in Guinea-Bissau
Data sources are provided at the end of thL country profile
J j._.i..^..
en
o m g
rsi
O)
T3
fti
(O
0)
.P
n n n
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('
Q)
a>
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t*
Q}
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m
u
03
t:
fr
E-
\r
t-
<
Tn
<
h
0)
CO
UJ
i^'^
<
o
u
z
Ed
^
363
'
World Atlas
Left:
Mangroves
in
Lagoas de Cufada
Natural Park.
Right: Forest destroyed
for cultivation.
Claudia Sousa
International Studies and
for
with
Cooperation ICECI]
from
support
financial
World
lUCN-Ifie
as duikers and other forest ungulates
is
also a
threat."
Conservation Union.'' More recent studies suggested their presence
in
the region of Xitole (an area
once proposed as a national park

Corubal
la
River], the
Ramsar
to the north of the
Lagoas de Cufada Natural
Parl<
the North Bank of Rio Grande de
Site),
Buba, Cantanhez Forest Hunting Reserve, and the

Cacine Basin.^''" The range
is
Boe
Guinea-Bissau
national Trade
119911,
119951.
sector,
in
the
southeastern regions of Quinara and Tombali.'
chimpanzee
habitat,
especially
primary
of
forest.'
Between 1990 and 2000, an estimated 220 kmforest
was
within
chimpanzee habitat
population
of
each year' Most land-use change
lost
increase.
constraints into account
ment Plan has
linked to local
is
Failure
in
to
human
take ecological
the National Develop-
also been identified as a leading
factor' Pressures include timber exploitation, bushfires,
clearing for agriculture, fruit farming, and
clearing of
mangroves
most common
in
for rice cultivation. This
is
becoming seriously fragmented by ba-
nana, cashew, and other plantations. Consequently,

crop-raiding by chimpanzees has also increased.
Chimpanzees are
not
generally eaten
humans. The young are sometimes taken
for
the local pet trade and chimpanzee skins are used

in
medicine.' Accidental capture of
traditional
chimpanzees
366
developing
in
snares set for
addition, all hunting
its
falls
protected area legislation.'
is
One
within the range of the western
chimpanzee: Lagoas de Cufada Natural Park, which
was
officially
declared
2000 and covers an area of
in
890 kml' Since 1990, about 44 percent of
this
park
has been internationally recognized as a Ramsar

Site."^
Possibly of
much
short-term survival
Bissau
in
is
resemblance
thought
greater significance for the
chimpanzees
of
in
Guinea-
the traditional protection afforded to
most parts
of the
them
country by their perceived close
to people.'
In
the Boe region they are
to shelter the spirits of elders."

Overview
There
is
game animals such
ample scope
measures
needed
in
Guinea-Bissau as they are considered too similar

to
In
is
the Tombali and Quinara regions.
The Cantanhez Forest Hunting Reserve iTombali

region)
Wild Fauna
hunting reserves. Guinea-Bissau
in
protected area
be the destruction
to
of
Ramsar Convention
and the Convention on Migratory Species
prohibited
still
The major threat seems
119901, the
under Decree No. 21/1980.

is
THREATS
Endangered Species
Chimpanzees are protected from hunting
between the
Corubal River and the Guinea border, and
in
and Flora ICITESl
Corubal
of the
has signed the Convention on
Biological Diversity 119951, the Convention on inter-
believed to extend
through the country to the south

River,' specifically in the
in
chimpanzee conservation
are:' to obtain basic information
anzee populations
feasibility
in
for
Guinea-Bissau. Three major actions
in
on chimp-
the country; to undertake a
study for establishing protected areas
the country and a transnational protected area
along
the
border between
Guinea; and
to
Guinea-Bissau and
develop a national strategy for
chimpanzee conservation.
AFRICA: Guinea-Bissau
Research and protected areas
size the
Future research should aim at developing action
tially
importance
of the primates, while
boosting the local economy.'
poten-
plans for chimpanzee conservation through studies
open woodlands, and
of their ecology, particularly in
through identifying appropriate protected areas for
two viable populations

Hunting Reserve and
of the Tombali,
Cantanhez Forest
at least: in
in
the Boe region.'
'
The basin
Cumbija, and Cacine Rivers, which
includes Cantanhez, has long been recognized as a
promising, high biodiversity area and

for protection.''
The status
'
of
the process of being upgraded
in
recommended
Cantanhez was
Capacity building
Long-term
between
collaboration
authorities, overseas
nongovernmental organizations
required
to
national system
of
and build national capacity
to
support the creation

protected
areas,
manage
effectively
it
government
governmental agencies, and
of
is
in
Education and tourism
2001.'
General education, awareness-raising programs,
and the involvement
Local participation
growing
Guinea-Bissau's
population
depends
heavily on forest resources. Biological conservation

is
therefore directly linked with economic grovsrth
and development. The participation
of rural
com-
munities by assuring them the legal right to
manage
at least part of the natural
could well be the best long-term

wildlife
resource base
way
to reconcile
conservation and rural development.
In
programs could help empha-
addition, ecotourism
conserving
can help greatly
habitat
local
of
chimpanzees and
to
communities
managing
relieve
in
their
pressure on
chimpanzees. There are currently no sanctuaries

for
orphaned chimpanzees. The establishment
rehabilitation center
and
its
resource would support the development
ape conservation
in
of a
use as an educational
of future
Guinea-Bissau. Alternatively,
confiscated animals could be sent to sanctuaries
in
neighboring countries.'
FURTHER READING
Gippoliti, S.,
Jones,
DelfOmo,
S. 11992)
G. 11996)
Guinea-Bissau.
Primates
In:
of the
Sayer,
J. A.,
Cantanhez forest and the Cacine basin, Guinea-Bissau. Oryx20: 74-80.

Harccurt, C.5., Collins, N.M., eds. The Conservation Atlas of Tropical
Forests: Africa. Macmlllan, London, pp. 200-205.
MAP DATA SOURCES

Map
16.12
following sources:
ButynskI, T.M. (20031 The chimpanzee Pan troglodytes: taxonomy, distribution, abundance, and conservation status.
Kormos,
R.,
Boesch, C, Bakarr,
M.I.,
ButynskI, T.M.. eds.
In:
Gippoliti, S.,
Dell'Omo, G. (20031 Primates of Guinea-Bissau, West Africa: distribution and conservation status. Primate
Conservation 19: 73-77.

Gippoliti, S.,
Embato,
D.S.,
Sousa,
C. (2003)
Guinea-Bissau.
In:
Kormos,
R.,
Boesch, C, Bakarr.
Plan.
M.I.,
ButynskI, TM., eds.
lUCN/SSC Primate
Specialist Group.
lUCN, Gland. Switzerland, pp. 55-61.

ACKNOWLEDGMENTS
This country study draws extensively on the Guinea-Bissau chapter from the
Status Survey and Conservation Action Plan.
Many thanks
to Brigid
lUCN/SSC West African Chimpanzees:
Barry [Tropical Biology Association] for editorial
assistance.
AUTHORS
Claudia Sousa,
Spartaco
New
Gippoliti,
Muhammad
University of Lisbon
Conservation Unit, PIstola Zoological Garden
Akhlas, University of East Anglia
365
World Atlas
REPUBLIC OF LIBERIA
Gemma Smith

Situated
West
in
republic, tiaving
Africa,
political,
Liberia
Africa's oldest
is
been established
1847 by freed
in
American slaves lAmerico-Liberiansl. Covering

area
total
of
km^
370
Leone
d'lvoire to the east. Sierra
Guinea
the largest city
is
The country has
in
to the
located on the coast
is
in Liberia.
and tablelands, and mount-
the far north. ^'
'^-
The
low/tands, with river-
and coastal vegetation, mangrove swamps,
ine
lagoons, and alluvial sandbars, are about 579
some
long and extend
65
country's agricultural land
km
is
inland.
Most
located
in
km
of the
the
hills
behind the lowlands. Plateaus and tablelands (up
300 m) and mountain ranges (up
to a height of
610 ml occur beyond
this area,
Lofa and Saint Paul Rivers

country.
mountain
in
in
mainly between the
the northwest of the
Highland areas, including the highest

Liberia (Mount Wuteve,
in
the north of the country,
counties.^"
currently
to
380 m) occur
Nimba and Lofa
in
An estimated 34 810 km^

under
still
of Liberia
is
forest: 31.3 percent of the total
upper Guinean
rain forest. Legal
and
illegal
logging are occurring rapidly, however, and forest
was estimated
cover
by the Food and Agriculture
Master Sergeant Samuel
then
of recurrent
calm, but
relatively
Doe overthrew
President William Tolbert after food price
riots.
Doe
being of the indigenous Krahn people, his coup
marked the departure from power
who had tended
country's establishment.
of instability
Americo-
of the
dominate since the
also heralded a period
It
human
as widespread
and
followed,
to
abuses
rights
provoked tensions between the
it
Krahn and other indigenous groups such as the

Mandingo,
and
and Mano." The economy collapsed
Gio,
all-out, ethnically
based
civil
war began
Dissidents of Charles Taylor's
INPFLI overran much
Forces of Liberia
NPFL
countryside, and an offshoot of the
of
the
killed
Doe
1990." These events prompted military inter-
in
Community
vention by the Economic
States
lECOWAS]
to protect
dents launched raids

in
1989.
in
National Patriotic
retaliation for
into Liberia
which the
West African
of
Monrovia. Liberian dissi-
from Sierra Leone,
NPFL supported
Sierra Leone, beginning Sierra Leone's

civil
war
rebels
own
in
10 year
(see Sierra Leone country profile].
The war
Liberia continued until 1996,
in
when
an ECOWAS-brokered peace agreement was signed,

eventually leading
president.
The
to
the election
Taylor as
of
conflict returned in 1999,
however,
Organization of the United Nations (FAOI to have
and escalated thereafter^ Under intense pressure
declined by 7.6 percent between 1990 and 2000.'
from
The
on Liberia (comprising the European Union, USA,
Liberia Forest
Reassessment
joint initiative of the
government
and Flora International

International
ICII,
1987 and 2001
project,
which
of Liberia,
is
Fauna
and Conservation
(FFI),
found total forest loss between
to
be only 2.6 percent, giving an
annual average forest loss rate
Africans, with Americo-Liberians
and
dants (colloquially referred to
Liberia as 'Congo'
after their
in
supposed geographical
their
descen-
origins)
com-
prising about 5 percent of the estimated population

of 3.32 million.
about
lives in
The annual population growth
1.7 percent.'
urban areas
The majority
in
rate
is
of the population
central Liberia.
the International Contact Group
ECOWAS
Morocco, UN,
Secretariat,
main factions came
the
Comprehensive Peace Agreement

exiling
Taylor to
in
Nations
Mission
October 2003, and
Liberia
built
up
began
to
in
a full strength of
15 000 peacekeeping troops by mid-2004.

in
peacekeeping,
justice,
addresses criminal
rights, child protection,
United
deploy
to
broad and robust mandate, and

it
2003, thereby
and creating a power-
Nigeria
in
and
sign
to
It
has a
addition
to
human
and public information, as
well as the environment and forestry.

This troubled
ting
consequences
modern
history has had devasta-
for Liberia's
economy, as
infra-
structure and social capital have been destroyed
is literate.'
Liberia has experienced intense
Nigeria,
Australia],
An estimated
57.5 percent of the adult population (over the age of

151
ECOWAS and
sharing National Transitional Government.
of 0.2 percent."
The country's people are mostly indigenous
366
U years
was
conflict. Until 1980, Liberia
land area. This includes the largest remaining portion of
1980. There has been a series of
in
conflicts since then, totaling
Liberians,
coastal lovi/lands, inland
flat
rolling hills, plateaus,
ains
northwest,
to the
and the Atlantic Ocean
to the north,
south. The capital Monrovia
and
bordered by Cote
is
it
and economic disruption since a
social,
coup
military
and sustained
and there has been
little
investment.
Few
figures
AFRICA:
Map
^
o
CM
in
a>
en
in
Liberia
this
Liberia
country profile
s
en
o
(1)
(U
,-
f*
f*
r'
Q.
ni
n)
(1)
Cl)
(1)
0)
fl>
Q]
(U
&
Q)
CD
OJ
(1)
tl)
fl)
05
(U
F
O}
CI)
5 5 ^
c
CD
ra
F
tn
UJ
Ir
367
World Atlas
are available, but the most recent 12002) estimates
serious
by the International Monetary Fund IIMFI suggest a
borders."
gross domestic product IGDPI of US$561.8 million
occurred
and
gross national income IGNII per person
of
US$169.20." There was insufficient information

available
in
2003 or 2004 for the United Nations
Development Programme lUNDP)
Human Development
country
is
certainly
an average
Index for
among
but the
the world's poorest, with
expectancy
life
Liberia,
of
only 41.7 years."
International aid has increased with the signing of
undertaken
WWF-The
by
was
Conservation
Global
the
Mano
River Union
and generated large numbers
Refugee populations are also known
in
south near the borders with Cote
Chimpanzees have been protected by law

Liberia since 1964, but the hunting of
for
in
chimpanzees
meat occurs throughout the country, and com-
mercial hunting
is
seen as
threat.""
a particular
The
rate of population decline,
the
Mount Nimba region hunted species such as
to the
chimpanzees
national survey of Liberia's wildlife
first
in
of 2003, factional fighting
d'lvoire.
Up
area,
in this
the east and
the Comprehensive Peace Agreement."
The
summer
the
In
of refugees.
calculate a
to
northern Liberia,
in
lifestyle.'
unknown.
is
Mano and
1960s, the
Gio peoples of
support their shifting cultivation
to
When
mining operations began
was heavy
there
area,
any,
if
exploitation
this
in
the
of
local
Organization and the country's Forest Development
wildlife (including
Authority IFDAI.^' Twelve primate species, including
workers.' Rural communities
the western chimpanzee [Pan troglodytes verus],
ceived chimpanzees as pests of their tree crops, and
live
in
Chimpanzees were probably once
Liberia.
present
in
forested
all
historically they
parts
Liberia," and
of
have been widely recorded
southeastern Liberian forests
in
the
in
Gola and
in
upper Nimba county along the borders with
Nimba Nature
the East
Grebo National Forest, Krahn-Bassa
Reserve,
National Forest, and Sapo National Park INP].''

In
1
the
1V70s,
000-5 000
it
was estimated
individuals.
that there
most chimpanzees were found
in
high forest
and older logged areas, where observations
of nut-
cracking sites and calls were relatively frequent.
local
population
THREATS
of forest in
east of the country,
from
in
the northwest and south-
particular, are facing intense
timber
and
extraction
operations.'' Subsistence agriculture
along roadsides, and around

in
The growing
hunting
commercial hunting, and pet-trade
pressures,
demand. Liberian bushmeat
is
marketed
the
in
Upper Guinea Forest subregion and may reach
global market. Internationally funded trade surveys
were being undertaken

confirm
in
2004,
an attempt
in
to
this.
some areas
In
of Liberia,
it
taboo
is
to eat
bushmeat, particularly within Islamic
primate
populations
in
the north of the country.
Nimba and among some
the southeast (for example the Sapo
and Kpanyan
districts of
eastern
In
ethnic groups and clans
Sinoe county],
it
Wehdjeh clan
in
Pynestown
in
is
taboo
to
Sapo
of the
(near the northern border of Sapo NP| consider
Large tracts
ments
continued
increases subsistence
eat chimpanzees. The
pressure
to the
military personnel, into rural areas.
were
Surveys carried out be-
tween December 1989 and March 1990 indicated

that
These pressures remain due
and logging industries, as well as military and non-
upper Lofa areas, and
in
so they hunted them for bushmeat and for the pet

trade.
the Sapo, Krahn
Cote d'lvoire and Guinea. Recent surveys have
confirmed their survival
mine-
migration of farmers and laborers from mining
northwestern and western forest
blocks of the
to feed the
the southeast per-
in
in
Bassa, Grebo, and Barrabo areas, as well as being
noted
chimpanzees!
new
is
mining
spreading
laborer settle-
has exacerbated threats
to
chimp-
to
be their relatives, from
are believed to have adapted
some
Consequently, they are forbidden
whom
they
forest skills.
to
kill
chim-
panzees." There are reports, however, that along

the
Liberia-Sierra
hunted
the forest.
Civil conflict
chimpanzees
Leone border the species
is
body parts that are used for medicinal
for
and magical purposes.
anzee populations, both through military operations
and the displacement

internally displaced
of
people.
Refugees and
persons are a defining element
of Liberia's post-conflict situation."
Nearly a million
people have been displaced - close to a third of the

country's population.
368
The
situation
is
particularly
LEGISLATION
Legislation
Liberia
is
party to the African Convention on the
Conservation of Nature and Natural Resources, the
in
Endangered
AFRICA:
Species of Wild Fauna and Flora (CITES), the Convention on Biological Diversity, the
Combat
to
Convention
the World
Desertification,
UN Frameworl<
Convention, the
UN
Heritage
Convention on
Climate Change, and the Convention on Wetlands
Importance IRamsarl. Liberia
of International
mid-200^, and has submitted
in
UN
National Report to the
Convention
first
its
Combat
to
Existing legislation,
and the associated
tutional framew/ork, should
There
Act,
of Liberia's
insti-
for the sustainable
allov\/
environmental resources.
an Environmental Protection Agency (EPAl
is
an Environmental Protection and Management

an Act Establishing a Protected Forest Areas
Lavi(,
and
Networl<,
National Environmental Policy.
managed
Liberia's forests are
velopment Authority
concessions
50 000 l<m^
by the Forest De-
has issued logging
vi/hich
The forest sector
legislation
has
While existing regulations appear appro-
priate, their
to
science." The area
enforcement has been weak.''
The
FFI
UK
government's Darwin
grant to
gazetted on April
management
laws and policies covering the
of
Liberia's environment.
Initiative
gave
of
September 2002. The
(GEFl
supported
and, as of mid-2004, so far the
only fully protected area
in
Liberia that had ever
for conservation."
has been the focus
of
much
servation effort. Originally
October 2003
preparation
management program
2005.
early
via
to
Created
in
983,
of the country's
it
con-
km^ comprises
swampy areas, and
it
be launched
FFI, to
Darwin grant and the beginning
of the
of a
GEF-
The Forest Development Authority and the

Society for the Conservation of Nature of Liberia
(SCNLl have carried out
field
resulted
basic
provision
in
of
These
activities.
equipment and
of
an 18 month operational plan, outreach
to
communities (environmental awareness,
provision of wells and latrines), and launching of a
A second Darwin Initiative

communal forests
bio-monitoring program.
was made
to FFI to pilot
NP
in
2004-2006. This
is
the forest resources that they traditionally
upon
for subsistence
uses, but
is
serving
communities
to
depend
and small-scale commercial
also establishing a buffer zone around
the park.
Since 1997, a
number
of
conservation projects have
been funded and implemented by FFI and
in Liberia.
in
supported program.
dryland
Sapo NP contains what may be the most
to several
Facility
long-term
Fund has provided bridging funding between the

end
lowland
forests.
ecosystem
073 km' but expanded
650
rain forest, including
2002, the
Ecosystem Partnership
Critical
Cl's
of
not only to secure the rights of rural

first
In
World Bank and the Global Environment
around Sapo
Sapo NP was the
the Whitley
Africa,
Foundation, and the Philadelphia Zoo.
grant
Protected areas
It
intact
remains reasonably
other forest areas to the north,
west, and southeast, extending into Cote d'lvoire.

is
of the park's
management
active
WWF-West
donors including
local
connected
restart
the park from April 2000 to
ment
forest
secure,
fully
infrastructure.'
November 26 2002 and

30 200^. The EPA Act contains
and riparian
to
new
infrastructure, allowances, basic training, develop-
created by legislature on
in
from
tar
is
however, as illustrated by the looting
The Environmental Protection Agency was
been managed
the western
in
were endemic
the Upper Guinean forest and six that were
30 companies covering more than
to
in total.'
been evaluated by the Liberia Forest Reassessment

project.
2002
late
in
funds were complemented by support from other
Desertification.
management
ten days
in
part of the park, including 78 that
and Action
lized its National Biodiversity Strategy
Plan
fina-
collected
It
thus at the heart of the largest remaining forest
Upper Guinean
latter
acting
Partnership
through
Fund and Centre
for Applied
diversity Science.
Both have worked
Liberian partners
in
forest
CI,
to
Bio-
support
restarting conservation
management, and
the
Ecosystem
Critical
its
and
build the capacity of
to
ecosystem,
several Liberian organizations, including the Forest
providing habitat to species that need to range over
Development Authority, the SCNL, the National
block
of
the
Liberia
forest
A faunal
large areas, such as forest elephants.
monitoring program, established

the park harbors
disturbed wildlife
some
in
in
2001 found that

,
of the richest
West
Africa's
and
rain
least
forests.
Chimpanzees have been surveyed," but botanical

surveys are less advanced: 353 plant species were
Environmental Commission
now
the
of Liberia
(NECOLIBI -
the Liberian Environmental Protection Agency,
Save
My Future Foundation, and Green
Advocates.
From 2001
to 2004, FFI
and
CI
implemented
the Liberia Forest Reassessment, a project that
369
World Atlas
aims
and
necessary information,
to establish the
policy
forest
tools,
environment for effective and sustainable
and biodiversity management
emphasizes correcting
in Liberia.''
It
a historical imbalance that
favored commercial use of forests over protecting
and factors that affect the supply
opinion,
bushmeat
line
market Isuch as the price
to the
of
gaso-
of
and ammunition]. Potential outlets through
which bushmeat enters international markets were

also studied.
representative samples of Liberia's biodiversity, and
meeting the economic and cultural needs

Liberians.
implemented
is
It
of rural
partnership with
in
three national agencies Ithe Forest Development
Department
Authority, the EPA, and the
of Statistics
Planning and Economic
of the Ministry of
Affairs],
Commis-
with financial support from the European
sion and CI's Critical Ecosystem Partnership Fund.
The aims
Reassessment
of the Liberia Forest
included an assessment of forest cover and the
management
protection status and
key forest
of
Sanctuaries
There are no sanctuaries that accept newly or-
phaned chimpanzees
oratory that
now
is
and
socialization
chimpanzees on
of
II,
a vaccine
is
the
lab-
slowly working towards the
release
of
ex-laboratory
its
small islands.
to six
chimpanzees had
VIlab
Liberia.
in
Hepatitis Research Foundation,
Ninety
been rehabilitated and
earlier
released,
but virtually all were
during the
civil
shot or starved
war
areas, with a view to updating Liberia's system of

legally protected forest areas.
Such areas may be
intended for forestry purposes, nature conservation,

research, or low-impact
human use
(for
as buffer zones and for recreation].

satellite
It
example,

Priorities for action
These include:
analyzed
imagery from the mid-1980s and 2001
to
comprehensive environmental assessment by
changes
of
government agencies supported by
reassess the extent
of,
Liberia's forest cover,
and quality
to,
and established a geographic
nongovernmental
information system (GIS] database that has been
assessment
continually expanded and updated with biophysical
its
and socioeconomic information. Liberian

policy
was
ment
provided, a
new system
area categories agreed, and
undertaken
ficant forest blocks
surveys
to obtain
classify
and manage
of
through
(V\/CF1,
is
CI's Critical
pro-
and protecting major populations
key species, including chimpanzees;
biodiversity surveys
ding
Between 2003 and 2006, the Wild Chimpanzee

Foundation
of
of existing
protected
areas ISapo. Nimbal;
to
diversity In the future.
Partnership Fund,
management
adequate management
adequate
and bio-
forests
Liberia's
biodiversity
of signi-
socioeconomic and biological data with which
and
tected area network representative of Liberia's
of forest protected
field
of the Liberian forestry sector
impacts on chimpanzee populations;
establishment and
forest
reviewed, recommendations for improve-
UN and
scientific organizations;^"
identification
and inventories, incluviable
of
populations of
chimpanzees; and
Ecosystem
implementing a U5$18A276
program
of
awareness].
environmental education (public
'"
education and awareness project to improve the

protection
of
wild
chimpanzees
in
West
activities include
performances and distribution
Africa.
dramatic
of newsletters,
and
building capacity so as to generate support from

local
people.
WCF
is
Liberian project partner
Parks Division
working
is
Development Authority
to July 200A, a
Concerned Environmentalists
of Biodiversity, In
Sapo NP. The
the Wildlife and National
of the Forest
From March
In
group called the
US$9 838
meat
370
mental administration
(albeit with a
forestry sector]. However,

conflict
and
is left
the
key
It
was
focus on the
looted during the
with almost no implementation or
enforcement capacity.
and
is
responsible for environ-
institution
It
needs
to
be re-equipped
re-skilled.
Enhancement
The Environmental Protection Agency also

requires strengthening and activation. Once this has
for the
project to analyze the Liberian bush-
trade. Surveys
obtain data on
government
partnership with Philadelphia Zoo
and the Bushmeat Crisis Task Force, implemented

a
Improved environmental governance
The Forest Development Authority
were conducted
in
Monrovia
to
volumes and species traded, public
occurred
It
supervise,
would be able
to coordinate, monitor,
and consult on
all
activities
In
the
protection of the environment and the sustainable
use
of natural resources.'
AFRICA: Mali
FURTHER READING
Agoramoorthy,
promise
Hsu, M.J. 119991 Rehabilitation and release of chimpanzees on a natural island - methods hold
G.,
for other
primates as well. Journal of Wildlife Rehabilitation 22
3-7.
111:
FFI In.d.l Liberia - Conservation in a Post-Conflict Country, http://www.fauna-flora.org/africa/feature_liberial.
html. Accessed June 30 2005.
UNEP
Desk Study on the Environment
120041
in
Liberia.
United Nations Environment Programme, Geneva.
http://postconflict.unep.ch/liberia/Liberia_DS_AGL.pdf. Accessed July 17 2004.
MAP DATA SOURCES

Map
16.13
Chimpanzee data are based on the following sources:
Butynski, T.M. 12003) The chimpanzee Pan troglodytes: taxonomy, distribution, abundance, and conservation status.
Kormos,
In:
R.,
Boesch, C, Bakarr,
M.I..
Butynski, T.M.. eds. West African Chimpanzees: Status Survey and
Nisbett, R.A., Peal, A.L., Hoyt, R.A., Carter,
eds.
J.
(20031 Liberia.
Kormos,
In:
R.,
Boesch, C, Bakarr, ML, Butynski, TM.,
Plan.
lUCN/SSC Primate
Specialist
Group. lUCN, Gland, Switzerland, pp. 89-98.

ACKNOWLEDGMENTS
This country study draws extensively on the Liberia chapter from the
Survey and Conservation Action Plan." Many thanks
Jamison Suter IFauna and Flora
COMPILER
Gemma Smith, UNEP
Chris Magin (Royal Society for the Protection of Birdsl and
to
comments on
International) for their valuable
World Conservation Monitoring Centre
REPUBLIC OF MALI
Chris Duvall and Gei>^ma Smith

The Republic
lying
1
central
in
241 138
which Bobo form the vast
of Mali is a large,
km^
West
Africa.
landlocked nation
It
bordered by Algeria
extends
Nigertotheeast, Mauritania, Senegal, and Guinea
to
the west, and Cote divoire and Burkina Paso to the
110 percent),
nial rule in
Modibo
overthrown
1968
in
in
a military
Moussa Traore.
uprising led by students
is
vast majority
live in
its
low for
relatively
population of about 12
its
large land area, the
the densely populated southern
third of the country.
The population
is
growing
2.8 percent per yean^ Mali's people are
nantly
Muslim but
ethnic groups.
include the
of
Mali's
predomi-
culturally diverse, with over 40
The most numerous ethnic groups
Mande
total
at
or Manding (about 50 percent
population,
comprising Bamanan,
of President
His single-party government
Kayes, Kidal, Koulikoro, Mopti, Segou, Sikasso, and
million^
percent),
16
1960 under the leadership
Keita.
south." Mali has eight administrative regions - Gao,
Tombouctou. Although
Touareg and
majority),
and Songhai
Mali gained independence from French colo-
over
the north,
to
Moor
Lieutenant Colonel
led by
in
popular
Bamako,
Amadou Toumani Toure
threw the Traore regime. Toure established

itional
government and
reforms leading
was
General
the capital
following
1991,
In
coup
democratic
instituted
to elections in 1992.
over-
a trans-
Alpha
Oumar
Konare, representing the Alliance for Democracy

Mali,
in
became
the
first
president and
was
in
re-elected
1997. Konare pursued a series of profound politi-
cal
reforms
to
democratize and decentralize Mall's
government and
Maninka, Soninke, and other groups), Peulh or
Mali
Fulani (17 percent). Voltaic groups (12 percent, of
ful,
is
now
privatize parts of
frequently cited as a
multi-party governance
its
economy, and
model
in Africa. In
of
success-
2002, Toure
371

World Atus
Map
16. 1i
Chimpanzee
distribution in Mali
country profile
this
CA--r-V--
Vxl
H
o
>
-^.
<>V^^'*i-
CO
CD
IT)
ro
g
D
(M
en
n n
T)
ro
0)
ai
(1)
(1)
r-
r*
i>
(1)
(i>
(1)
i>
n)
Q-
ai
(U
Q)
a)
(1)
a
b
n
b
n
b
n
E
<ti
ai
0)
0)
a Z
a)
(11
<:
fH
s s s
ra
lU
Lr
'<-
^l'
cj
372
* L
t\'
AFRICA: Mali
elected as an independent candidate to replace
was
main tributaries - the Bating, Bakoye, Baoule, and
Konare." Toures domestic policies have focused
Faleme Rivers
on the struggling econonny, Mali's unemployed
plains
and poorly educated youth, and the
the involvement of local
nongovern-
society,
civil
mental organizations INGOsI, and other stakeholders
government and economic development,
in
women.
the worlds poorest
with a growing emphasis on involving
remains one
Mali
of
Gross domestic product IGDP)
nations.
200/1
in
none
More than two
tight against
corruption. Decentralization continues to increase
arid,
to
which have wide flood-
thirds of Mall
Is
and or semi-
with vegetation density increasing from north
south as annual rainfall increases from less than
mm
100
in
of
in Mali."'
more than
the Sahara to
In
the south.'' The country
mm
500
divided latitudlnally
is
"''
three distinct zones of natural vegetation:''
into
the Saharan zone (dominated by desert and semldesertl, the Sahelian
zone (dominated by Acacia
gross national income IGNII of about US$290 per
wooded grassland and deciduous shrublandl,

and the Sudano-Guinean woodland zone. The
person.^^ Although large areas of the north are
Sudano-Gulnean woodland zone covers about
was estimated
US$A.3
at
barren desert, the
representing a
billion,
Niger River basin and
fertile
km^
131 860
of Mali, 10.8 percent of Its land area.'"
dense
other parts of southern Mali support subsistence
These woodlands, which have
agriculture and animal husbandry, which have long
tree cover
dominated the economy. Manufacturing and indus-
Combretaceae and Leguminosae, are prominent
trial
gold mining
greatly
southwest Mali have increased
in
the past decade, and cotton, gold, and
in
a relatively
dominated by various species
the landscape
the
in
in
areas where chimpanzees occur
in
Mall's closed-canopy forests are mainly located
in
livestock are the nation's
most valuable exports.
the southwest, and are primarily riparian gallery
Interannual variability
weather has
forests
in
strong
influence on agricultural production and therefore
GDP.
In
foreign
2002,
about
contributed
aid
percent of GDP," and Mali has recently benefited
from debt
off
by France
relief
including a 38 percent debt write-
relief,
2002, and
in
US$765
million
debt
in
from the International Monetary Fund lIMFI
and the World Bank
in
2003. Mall's debt burden
government's
remains high, and
limits the
to Invest in sectors
such as education, healthcare,
ability
dominated by bamboo [Oxytenanthera

Graminae) and raphia palm {Raphia
abyssinica,
Palmael.' Additionally, small patches of non-
sp.,
riparian gallery forest occur on steep slopes and
edges
cliff
In
dominates these
types
grassland and shrubland, with few trees
of
areas with shallow or poor
development.
southern Mali.
Is
mostly
flat
lowest elevation
Senegal
River],
Is
23
200-500 m.^^ The
above sea
of
is
155
level
Ithe
IHombori
southern Mall
lies in
the Niger River floodplain, with the only relief being
small areas
river's
undulating
of
various tributaries.
In
large patches
throughout
soil
and low-lying, with
of
and the highest
Tondo rock outcrop). Most
and many species with
and dominated by grasses, occur

In
Mali's terrain
forests,
edible fruits are also abundant.' Finally, several
natural resource conservation, and infrastructure
an average altltudlnal range
the Manding Plateau. The endemic
Giltetiodendron gtanduiosum (Leguminosae!
tree
between the
hills lying
However,
In
western,

Western chimpanzees [Pan troglodytes verus] are
the only species of great ape
reported by Sayer'
occur
" " They
In
Mall,
as
first
known
are only
to
the southern part of the western region of
In
the country.
the Manding Plateau near the bor-
In
ders of Guinea and Senegal.'
'
Most
of
southwest
central, and eastern Mali, sandstone massifs out-
Mali has not been surveyed tor chimpanzees, but
crop to form topographically complex highland
isestimated that the
areas that have a relatively high level

diversity
due
to the
wide range
of
supported by the varied topography.'

areas include; the Dogon
Cliffs in
Adrar des Iforhas, located

northeast;
and
in
in
of biological
microhabitats
"
''
These
central Mali; the
the Sahara
the southwest,
the
in
the
Manding
where chimpanzees occur in

including most of the Manding
Plateau, the only area

Mall.
Western
Plateau,
is
Mali,
drained by the Senegal River and
Its
species
in
Mall
Is
maximum
19 440
kml'
it
current range of the

Mali's
chimpanzees
are the northernmost natural population, occurring

to
about 1310'N, Including
environments
in
some
of
the driest
the animal's global range."'"
In
the
recent past, chimpanzees were found farther north
than at present;" this range decline
due
to
known
human
if
is
threats, especially hunting.'
probably
It
the Malian chimpanzee population
rently isolated
from those
in
is
Is
not
cur-
neighboring countries.
373
"
World Atlas
shows
but genetic analysis

isolated
Chimpanzees
been
gallery forest
Mali,
in
riparian
non-riparian
particularly
in
cliffs
and
the
in
(^/landing
to
be an exception
known
little is
to
The
for certain.'
in
mam
and agricultural expansion, although Mall's chim-
'^
" and evidence
chimpanzee
of
as feeding remains) has been found
in
There have been two efforts
chimpanzee population density
estimate
to
on nest sampling. Pavy calculated
chimpanzees per square kilometer
based
density of
the nor-
in
thern portion of the proposed Bating Biosphere
Granier and
while
panzees appear
Martinez
found
Mall are hunting
In
be relatively less vulnerable
to
to
these threats than most other natural populations.''
Chimpanzees are
to
Mali, both
in
threats to chimpanzees
and
bare sandstone with minimal shrub cover'
other large
rarely
mammal
low population size
In
hunted
'^
species,'
Mali
absolute levels of hunting
Mall relative
In
but the animal's
means that even low

may pose a relatively
problem." Malians most frequently hunt
large
chimpanzees with guns
reduce losses
to
of fruit
from wild trees, although baboons and monkeys

are the primary target species.
the parts of Mali
In
evidence of 0.35-0.40 chimpanzees per square
where chimpanzees occur, Indigenous
kilometer
beliefs that have no restrictions on eating primate
border,
among
three limited areas near the Guinea
in
south,
and west
east,
Both
reserve.'^
of
proposed
the
these density estimates are
of
the highest recorded for chimpanzees
tiously as the
density
Mall.
in
upper
likely
limit
of
chimpanzee
Recognizing that Granier and
Martinez's estimate applies only to the
limited
areas where they collected data, Pavys estimate
more
likely to
represent the
maximum
in
population
based on the range area cited above,
size for Mall,

is
maximum chimpanzee
Is
average
population density for the animal's entire range

Mall. Thus, the
in
must be viewed cau-
similar habitats,'^ and thus
h 860 individuals, although the actual population
size
is
certainly
lower,
patchlly distributed.^'
since
chimpanzees are
'^
and widespread
south
least densely
of the
only
the west of the Bating River,

is
one
of the
populated parts of West Africa south
Sahara. On the west bank of the Bating
lie
National Parks (NPs) and
Chimpanzee Sanctuary
Bating
specific
Wongo
(a
species-
reserve for naturally occurring
faunal
many consider chimpanzees

humans to want to eat them.'
but
to
proposed
to
Mall has been undertaken
In
An
In
the
additional protected area has been
the
south
of
the
proposed Bating
rural areas do not recognize, whilst
in
chimpanzees as
aware
to
of
Mali and
Guinea
in
an isolated and biologically
to
them
many are
Further potential threat
Itself.'
willing
an opportunity
If
Is
posed
by hunters from nations to the south, particularly

Guinea, where chimpanzee meat
consumed, who may extend
more widely
Is
their hunting activities
northwards.
Habitat loss
to
Is
Mall.'
medlum-
probably the greatest
long-term threat
to
chimpanzee
survival
In
and has four main causes: farming, livestock
herding, logging, and mining. Although chimpan-
zees are most abundant
is
cliffs
in
areas that are not
and steep
a threat in
some
agricultural
hills,
areas,''
" particularly
east of the Bating River and south of the town of
Manantall. Traditional swidden
(slash-and-burnl
farming requires large land areas, but
human
diverse area."
Rural Mallan hunters are
pets.
these urban markets, and
supply chimpanzees
Faleme Resen/e would
in
some
wealthy Malians and European expatriates value
term
conservation efforts
supply the
medicinal properties or spiritual significance that
people
Biosphere Reserve; the transboundary Bafinglink
less frequent
to
is
consider chimpanzee skin and other parts to have
expansion
Bating area.
and young chimpanzees. Many urban Malians
Bating Biosphere Reserve.' Nearly
chimpanzees
be too similar
to
small, largely urban market for skins, body parts,
farmed, such as
research on
chimpanzee
eat
'^
reason for hunting chimpanzees
populations!, which together comprise the proposed

all
religious
meat, either as food or for medicinal reasons,
the
Mallan protected areas with chimpanzees:
Kcuroufing and
the
to
Manantall Dam,' " which
of the
meat predominate. Some people do
presents
Chimpanzees are apparently more abundant
376
though
Mali
at
a wide range of habitats, including grasslands
Reserve, ''
and chimpanzees are unlikely

this pattern,
in
the last 50 years,^'^'"
In
away from water sources,
least seasonally,'
0.25
most large mammals
of
have declined markedly
Chinnpanzees also nest and feed
situations,
activity (such
THREATS
The populations
inhabit areas of
gallery forest patches

Plateau.''^ "^'
woodland
that they have not
the past."
in
Is
sustainable approach
In
In
the long
areas
of
low
population density. Population growth and
increased market orientation are changing agricultural practices,'
and permanent loss
of
woodland
AFRICA: Mali
areas
to agriculture
increasing around larger
is
settlements and along all-season roads." Since
about 1989. following the construction

across
tfie
Bating
River at
bridge
of a
Manantali Dam.
ttie
livestock herders from northern Mali have entered
proposed Bating reserve each dry season.
the
Herders
fell
provide forage for their
trees to
animals, rather than relying solely on herbaceous

vegetation. While the effects of their activities on
wildlife
have not been studied, local residents
blame the herders

wildfires
the herders do
increased incidence
for
visit
Chimpanzee
of
habitat in
abundance, and
the proposed Bating
areas and cut trees used by
Biosphere Reserve.
and a decrease
in wildlife
chimpanzees.' Small-scale logging has also re-
duced chimpanzee habitat since the

Manantali
Reservoir
filling of
the
Fishermen based
1988.
in
along the Manantali Reservoir (including within the
proposed reserve!
illegally
harvest large hardwood
species for canoe-building. creating major canopy
gaps
in
riparian forests.' Industrial mining oper-
ations near Kenieba have cleared vegetation and

soil
from areas
habitat,
of
l<nown or potential chimpanzee
produced much
and triggered an
and nontoxic waste,
toxic
of jobseekers.'^
influx
Finally,
proposals to build a major international highway

just south of the
proposed Bating reserve
funded, probably increase

in
the future. Habitat loss
all
is
forms
a
if
of habitat loss
more
sure to address than hunting, as
will,
pres-
difficult
results from
it
the economically important activities of farming,

livestock herding, fishing,
areas have been designated
protected
about 5 percent
covering
newest
of
Mali,
in
country.''
The
these are Kouroufing and
Wongo NPs
2001), and the Bating
Chimpanzee
(both gazetted
in
Sanctuary (gazetted
in
An extensive
20021.
zone and a zone cynegetique
buffer
multipurpose area
(a
intended to protect defaunated woodland habitat
whose
exact status has not been determined! are
proposed additions
to
the three gazetted areas.
Altogether these protected areas form the proposed

Bating Biosphere Reserve, a total area of about
5 215 km'.'
drawn up
in
A management
plan for this reserve
was
2000-2001, but as yet only limited funds
have been allocated for
its
implementation. First
established as a faunal reserve
1990, the Bating
in
Chimpanzee Sanctuary remains poorly developed.

Since the early 1980s, chimpanzee protection has
and mining.
been one

Mali has signed and ratified a
the
of
number
of
primary goals
the
As
activities in the area.
of inter-
for the
of
conservation
proposed Bafing-
area, the Direction Nationale de
Faleme protected
Conservation de
Nature
actively undertaking
national biodiversity and conservation conventions
la
and agreements. These include the Convention on
natural resource assessments of the area and
Biological Diversity, the Convention on
Wetlands
of
likely to
national Trade
in
Endangered Species
of
Wild Fauna
and Flora (CITES), the World Heritage Convention,

the
the
UN Convention
UN Framework
to
Combat
Desertification,
and
Convention on Climate Change.
it
responsibility of the
Conservation de
in
1998
to take
la
is
of wildlife
Direction
Nature,' which
over
many
in
Mali
governs wildlife management, and are
Appendix
Conservation de
Nature (DNCNI has
la
several planning and
management
initiated
projects
in
the
Protected areas are designated under

No. 86-43/AN-RM,
86-^2/AN-RM
of that law,
Law
listed
in
which gives them complete
protection against capture, habitat destruction, and

hunting.'
The minister responsible

la
Conservation de
la
for the Direction
Nature can grant
exceptions to this rule for scientific collection and
concerning hunting and the
research, or the removal of dangerous animals.
and under Law
Other Malian laws, such as that governing forest
conservation of fauna and
No.
Nationale de
proposed Bating Biosphere Reserve.
Chimpan-
zees are protected under Law No. 95-031, which
la
obligations under one or
la
functions of the former
Direction Nationale des Eaux et Forets.
of
its
the
was established
more
of
is
Nationale de
these agreements, the Direction Nationale de
As part
is
the near future.
in
The management
International Importance IRamsarl, the Convention
on Migratory Species, the Convention on Inter-
gazette
la
(the
its
habitat,
Forest Code). Thirteen
management (Law
No. 95-OOAl, provide protection
375
World Atlas
chimpanzee
for
particularly riparian
habitat,
and
members
figured
gallery forests, although the relevant parts of this
condemned them
law are rarely enforced.'"
unclothed
hunting and most woodcutting were illegal
All
prior to the restoration of civilian rule in 1992, after
which natural resource laws were modified
to suit
the democratic, decentralized principles of Konare's
reform
program.'
Current
management laws
and
wildlife
forest
some communityalthough how this de-
allow for
based resource control,
volution of authority will be undertal<en remains
The
unclear.
over
authority
decisions
made
state
in
management
resource
natural
the Bating area, including the decision
2001-2002
in
continues to exercise strong
to forcibly
remove several small
refused
chimpanzee clan and

raw food and remaining
of
the bush because the chimpanzees
in
to
the
to eating
abandon
a settlement site coveted by the
sorcerer' This story suggests that on a cultural
Maninka recognize that humans and chimp-
level
anzees compete
for natural resources,
many Maninka
by
nized
as recog-
and
individuals'
by
researchers.''''^^
Within chimpanzee range

lor Fula, Peulh, or Fulfulde) are
southern part
of the
Mali, the Fulani
in
mostly found
the
in
Manding Plateau, where they
practice sedentary farming. Fulani to the north of
the plateau practice pastoral livestock husbandry,
and
chimpanzee range only
visit
in
the dry season.
settlements from within the protected area. Local
For the most part, Fulani and Maninka attitudes
residents did not support this decision, but from the
toward chimpanzees are similar, although Fulani
move
standpoint of wildlife conservation, the
probably reduce hunting and habitat loss.

also increase
human
activities
along the
It
will
may
gazetted protected areas, thus perhaps negatively
chimpanzees.
Nationale de
la
general, the
In
Conservation de
la
anzee range areas. Many

migrants from elsewhere
been designated
that have
in
the chimp-
in
residents and
local
some
throughout the year, and
Direction
Nature has never
had a large or highly active presence
hunt actively
Mali
farm
still
eat
in
areas
in
southwest Mali are generally less
Urban Malians often have
different views
and
values concerning chimpanzees from rural people.
many urban people do not know what chimpare, and do not know the Maninka or
Bamanan names for the animal. Second, few of
those who do know of chimpanzees are indifferent
to them - many urban Malians consider them to be
ferocious and highly dangerous. Many people enjoy
First,
anzees
stories of the attacks or depredations of various

wildlife species, including
off-limits.
likely to
chimpanzee meat.'^
cliffs that
are located predominantly outside of the currently
affecting
people
chimpanzees, although
very few urban Malians have ever seen a chimp-
TRADITIONS
Two
anzee, and even fewer have seen a noncaptive
ethnic groups, the Maninl<a and the Fulani
dominate
found.'
areas where chimpanzees are
the
in
Maninka are also numerous
chimpanzee range
divoire, while Fulani also coexist with

in
parts of
in
Senegal, Guinea, and Cote
in
skin of
Finally,
for the spiritual
in
(or
the Manding
Plateau. Most rural Maninka are indifferent toward
chimpanzees,'^
although
animal range from fear
toward
attitudes
to respect,
and from
the
dislike
to focus
believed to add to certain
to
humansl.
sympathy Idue
Some
anzees a pest
rural
to their
resemblance
Maninka consider chimp-
of fruit trees
and
millet,^'
''
and many
consider the animal valuable, since consumption
meat
is
believed to treat
lonchocerciasis).^
known by some elders
blindness
is
hindered by
in
Two research
2003-2004 may improve the
projects
efficacy of
future conservation actions regarding the protection
chimpanzees. Researchers working with the

la
Gestion Integree des Ressources
Naturelles (AGIRI assessed

bution
chimpanzee
distri-
and population density, and threats
chimpanzees
at several sites in
southwest Mali
to
in
2003-2004.'^ Their findings provide baseline data
Maninka story
for the evaluation of future conservation efforts, as
the Bating area tells that
well as improved understanding of the international
traditional
in
river
Mali
in
conservation efforts
conducted
project Appui a
human
significance
of
Mali's
the distant past, a powerful sorcerer dis-
assessment
of
chimpanzee
chimpanzees represent one

In
is
lack of resources and information on the best sites
of
clans.
it
Chimpanzee conservation
humansl
its
power
people value the
the fabrication of amulets
prayers or charms.'
(due to their consumption of wild foods valued by
of
in
Malinkel are the main ethnic group
to
many urban
chimpanzees
chimpanzees
Guinea.
Sedentary agriculturalists, the Maninl<a
376
chimpanzee.
of
several
chimpanzees. Second, an
habitat
and threats
AFRICA: Mali
posed by subsistence farming and hunting

Bating area
was undertaken
recommended
the
in
2003-2004 with the
in
Great Ape Consen/ation
of improving effective management. The

same recommendation should be made for the
Conservation
Fund,
proposed Bafing-Faleme transboundary protected
and the Milwaukee Zoological
International
(Cll,
Society.' This
research will help clarify ecological
area. Sustainable use of natural resources
and spatial relationships between agriculture and
human
settlement,
areas should be allowed as specified
and chimpanzee distribution
legislation,
and abundance.
to
West African Chimpanzees: Status Survey
In

colleagues
focal
recommended
Mali.'
in
in
Mali
were
posed Bating Biosphere Reserve, the area

east of the Bating River, the
Tambaoura
ed
to the
the
Tambaoura
publication
Cliffs
Second,
plan.
Conservation de
la
these
Association
Faune
Finally, efforts are
et
de
need-
increase public awareness of the international
chimpanzee population, and
toward chimpanzees.'^
activities
such as those
begun by the Wild Chimpanzee Foundation (WCFI
was
it
to
NGO
the
to dispel negative attitudes
have been studied since the
the action
of
la
significance of Mali's
but
all
effectively enforced
for local residents in
by
lEnvironnement lAMCFEI.
Keniebal, and along the Faleme River on the
Mali-Senegal border Of these four areas,
pioneered
Malienne pour
Cliffs (north
these
introduce and stimulate alternative de-
as
areas,
identified: the pro-
in
the relevant
protected areas. Concerted efforts are
in
to
velopment opportunities
four
First,
areas for chimpanzee conservation and/or
further research
of
needed
several conservation
chimpanzees
for
priorities
which must be more
in
reduce poaching, livestock herding, logging, and
farming
Chris Duvall and
Plan,
the
to
aim
support of the Wildlife Conservation Society IWCSI,

the
support
financial
that
proposed Bating reserve be increased with the
must be continued and further expanded.
FURTHER READING
Duvall, C. (2003) Agriculture
Butynski, T.M., eds,
and chimpanzee survival
West
Africa. In:
Kormos,
Boesch, C, Bakarr, ML,
R.,
West African Cfiimpanzees: Status Survey and Conservation Action
Plan. lUCN, Gland.
Switzerland, pp. 143-145.

Duvall, C. (2000) Important habitat for
Moore,
J.
Chimpanzee survey
(1985)
in
chimpanzees
Mali,
West
in
Mali. African
Africa.
Deuxieme rapport du Mali sur
Toure, A.S.. MaVga, A. (2001)
Study Monograpfis2\
Primate Conservation
6:
141:
173-203.
59-63.
la diversite biologique.
http://www.biodiv.org/doc/
wcrld/ml/ml-nr-02-frdoc. Accessed November 5 2004.
MAP DATA SOURCES

Map
16.14
Chimpanzee data are based on the following sources, with additional data
from Duvall,

status.
In:
by personal
communication
C. (2005):
Kormos,
R.,
Pan
troglodytes: taxonomy, distribution, abundance, and conservation
Boesch, C, Bakarr,
M.I.,
Butynski, T.M., eds. West African Chimpanzees: Status
Survey and Conservation Action Plan. lUCN/SSC Primate Specialist Group. lUCN. Gland, Switzerland.
pp. 5-12.
C, Niagate,
Duvall,
B.,
Pavy, J-M. (2003) Mali.
African Chimpanzees: Status Survey
In:
Kormos,
R.,
Boesch, C, Bakarr,
Plan.
M.I.,
Butynski, TM., eds. West
lUCN/SSC Primate
Specialist Group. lUCN,
Gland, Switzerland, pp. 41-50.

ACKNOWLEDGMENTS
This country study draws extensively on the Mali chapter from the
Survey and Conservation Action Plan. Many thanks

on the draft
of this section,
and
to
Jen Duvall for
to Ian
all
Redmond
(Ape Alliance/GRASP) tor his valuable
comments
her support.
AUTHORS
Chris Duvall, University of Wisconsin-Madison
Gemma
Smith,
UNEP World
377
World Atlas
FEDERAL REPUBLIC OF NIGERIA

Edmund McManus

The Federal Republic
damaged
country bordered by Benin,
Cameroon,
923 768
covering
km"
It
Chad, and
Niger,
area
surface
land
of
most populous country,
Africa's
Is
West African
of Nigeria is a
owed about US$28
country, which
billion
external creditors. Development has historically
been hindered by pervasive corruption, which the

Nigerian government
tackle, as well as
number
taking a
is
working
to
of steps to
improve relations with
with about 134 million people (estimates range
the International Monetary Fund (IMF], World Bank,
from 128
and Paris Club
137 million)" and an annual population
to
growth rate
independent
of
about 2.5 percent." Nigeria became

1960 as a federation of
of Britain in
sub-Saharan
leading
three regions INorthern, Western, and Eastern],
barrels of
investment
1963, Nigeria
became
self-government.'
of
a Federal Republic, with a
new
fourth region (the Midwest], and a
January 1966,
In
group
of
per day' Offshore
oil
The gross domestic product (GDPl

US$35.1
or US$1
billion,'
officers
overthrew the government and imposed a
purchasing power
state of
emergency Anti-lgbo
the
3.7 percent'
north,
and there was a counter-coup by northern
comes from
troops
in July.
in
Thousands
riots
broke out
in
Igbos were massacred
of
the north, and hundreds of thousands fled to
homelands
their
were increasingly strong
new
calls for secession.
The
military leadership replaced the regions with
12 states
in
an attempt
increase local self-
to
governance, but this gesture was rejected by the

military
government
Colonel
Eastern Region, under
of the
Odumegwu Ojukwu, who
declared that
region to be an independent state, the Republic of

Biafra'. Civil
when
war
followed, and lasted until 1970,
the Biafran troops surrendered.
The
970s,
980s, and
per person
to
parity,'
in
2003 was
adjusted for
billion
and was growing
oil
sales.' Nevertheless,
Nigeria
in
low, at
is
now
at
GDP
per year. About 20 percent of
Nigeria
annual income
US$3U,' or US$900
parity.'
consists of 36 states, plus the
Federal Capital Territory of Abuja. The climate

equatorial
in
the south, tropical
arid in the north.
lands
in
The
the south to
in
terrain varies
hills
southeast (extensions
of
from the low-
and plateaus
the
the central
in
in
in
the
Cameroon Highlands]
to plains in the north. Nigerian
from tropical rainforest
is
the center, and
region (Jos Plateau], and from mountains
in
990s saw a succession
U.8
adjusted for purchasing power
where there
the southeast,
in
production and
oil
gas sector are expected
the
in
the
is
producer at 2.3 million
oil
continue growing rapidly.
constitution.
mostly Igbo army
The economy has
of creditor nations.
considerable potential to recover, as Nigeria
which retained a large measure

In
ecosystems range
the south, dry thorn scrub
montane communities on the
the northeast, and
Mambilla Plateau and Obudu highlands near the
of military coups, with long periods of military rule
border with Cameroon, to the mangrove and fresh-
and intervals
water
government.
of civilian
The most recent

Sani Abacha from
dictatorship, that of General
1993
1998, dismantled all
to
swamp
swamp
forests of the
forests
were
and west by lowland
Niger Delta. These
originally flanked to the east

rain forest,
much
of
which has
democratic structures. This regime oversaw the
now been
execution of Ken Saro Wiwa, the Ogoni activist
largely subsistence-based agricultural sector has
and poet, and the systematic misappropriation

oil
to
revenues. General Abacha alone
have stolen US$A.3
billion.'
after General Abacha's
successors repealed
death
many
planned the restoration
food exporter, Nigeria
Nigeria
was sus-
remaining areas
in
until
1998. Abacha's
democratic,
government. Local elections were held
in
999,
and was won by
the southeast,
civilian
Decemin
former president,
Olusegun Obasanjo.
President Obasanjo inherited an economically
of
in
now imports food. The
largest
closed-canopy rain forest are
Cross River
state.
in
These are
contiguous with the forests of southwest Cameroon.
and
ber 1998; the presidential election followed
February
not kept up with population growth: once a large
estimated
military decrees
of a
of
replaced by cultivated land. However, the
is
pended from the Commonwealth from 1995
378
to
There are no great apes

part of the country.
In
in
the northernmost
the regions inhabited by
great apes, annual precipitation ranges from

to U
000
mm,
with a three to five
from November
people
live
to
500
month dry season
March." About
half of Nigeria's
within the historical range of chimpan-
zees, across the southern part of the country.
AFRICA: Nigeria
Map
in
Nigeria
ttiis
country profile
V3a
379
World Atlas
largest remaining
population
Gashaka Gumti NP and
large population occurring
and
probably that
is
vicinity,
its
Wildlife Sanctuary.''
NP
the Cross River
in
surroundings, including the
its
in
with another
Mountain
Afi
The chimpanzee range can be
main regions.
divided into three
Southwest Nigeria and the Niger Delta

Surveys are few
western Nigeria and along the
in
eastern edge of the Niger Delta. Chimpanzee populations here are small, highly fragmented, and
severely threatened." They
confirmed
to survive in the
the Niger Delta, and
Women from
village on the
Old Ekuri, a
edge
Omo,
in
have recently been
southeastern forests of
Ise,
Owo, and Okomu
Forest Resen/es. They are reported but not con-
firmed from the Oba
of
Hills, Ala,
Idanre, Ifon,
Okomu
Onishere, and
Ohusu
Ogbesse Forest Reserves,' and suspected
Park, collecting a forest
NP and
vegetable called afang
Forest Reserves.
the Akure-Ofusu,
and
in
iGnetum africanum).
Tunde Morakinyo

By 1982,
extinct
that
in
it
was
believed that gorillas had
become
Nigeria," but later surveys demonstrated
they are
still
These Cross River
present.
gorillas are recorded in the Afi
Sanctuary
Mountain Wildlife
River Forest
the Afi
of
Reserve,
in
Mbe Mountains community forest, and in the

Okwangwo Division of the Cross River National
the
Park INPl." There are probably three
distinct
to
individuals remaining
in
There
is
some
be approximately 80-100
Nigeria."
uncertainty about the sub-
chimpanzees found
the
specific
affinities
Nigeria.
Based on sequencing
of
DNA, chimpanzee populations

are related to populations
of
in
eastern Nigeria
but are not closely related to populations
Upper Guinea region or

the
in
Sanaga River
the
in
mitochondrial
western Cameroon,
in
in
the
chimpanzees south
to
border,
to
the Gashaka Gumti
in
Taraba and
in
Adamawa
NP
in
the country
on the eastern
states.
It
hosts up
500 chimpanzees. Chimpanzees also occur
in
neighboring areas, including the Mambilla Plateau

(specifically
in
Ngel Nyaki Forest Reserve and
Leinde Fadali and Akwaizantar forests! and the
Donga River valley"
of
Cross River state

There are thought
to
be around 400 chimpanzees
living in the
Okwangwo
River NP,
the northern part of Cross River state.
in
This population
the adjacent
is
Division of Nigeria's Cross
contiguous with populations
Takamanda
in
Forest Reserve of south-
west Cameroon and the Mbe Mountains Community

Forest
still
Nigeria.
in
The Mbe population
tenuously linked
Wildlife
to that in
is
probably
the Afi Mountain
Sanctuary and other areas
of the Afi River
Forest Reserve immediately to the west of Mbe,
Nigeria-Cameroon border region have
although the Ikom-Obudu highway runs between
troglodytes
in
vellerosus'-
analytical techniques used,
'
the
Depending
Reserve."
chimpanzees
P.
t.
on
in
the
west-
vellerosus as
shown on Map 16.15, or with P.

verus of Upper
Guinea. More research is needed, therefore, to
(.
the
affinities
of
chimpanzees
in
south-
western Nigeria.
A 2005 workshop
Mbe Mountains and
the subspecies Pan
ern Nigeria group either with
the
Oban
In
in
Brazzaville estimated that

in
Nigeria.
The
the
Afi
River
Forest
the southern part of Cross River state,
ing forest reserves
NP and
surround-
and community forests are also
important for chimpanzees.
THREATS
The major threats
there are around 3 050 chimpanzees
380
found
Cameroon. The chimpanzees
in
accordingly been placed
clarify
is
sub-
populations, and a fourth shared wîth Cameroon.
There are estimated
Gashaka-Mambilla
The largest chimpanzee population
survival
in
to
gorilla
and chimpanzee
Nigeria are hunting, forest degradation,
and deforestation.
In
2000,
it
was estimated
that
AFRICA: Nigeria
km
135 170
remained
forest
of
an average annual decrease
km^ or
3 980
cessions
in
almost
all forest
reserves
protected land area had al-
of Nigeria's
plantations,
of agriculture, oil
and road networks has
led to the
widespread degradation and fragmentation
of great
Deepening poverty
gap between
and an increased
levels
and poor, accompanied by a
rich
growing awareness
of
wealthy
combine
lifestyles,
to increase rural people's incentives to exploit forest
rities is
The relationship with national park autho-
sometimes
when
building
a difficult one, with
resentment
such
products
forest
as
plant
material or bushmeat are confiscated.
Chimpanzee meat
most
of
may
exist.
is
There
in
is
less hunting of
chimpanzees
nevertheless reported there."
in
in
is
unlikely that any
is
It
populations are entirely free from hunting pressure,

it
more
is
of
often opportunistic than planned.
sometimes
these infants
sold as pets, and the value
may encourage
hunters to target
Hunting has historicallythreatened the survival
Cross River
gorillas. In
many were
twice as
naira
989,
killed
born.' At that time
was 150
it
was suggested
that
each year as were being
reasonable monthly wage
per month, and
carcass could fetch as
much
15 communities hunted
in
1986 just one
single
gorilla
as 300 naira. About
Cameroon border
reduced,
at
Ekang, and there
is
also a
plan to improve the transborder highway from Ikom

to Ivlamfe.
LEGISUTION AND CONSERVATION ACTION

Legislation
Nigeria
Nature and Natural Resources
of
Endangered Species
(CITES)
in
Diversity
1985
Convention on the
the African
ratified
the
1968,
of
Fauna and Flora
Wild
1996.
in
The Endangered Species Act
hunting
in
is
in
protected by law, but these laws

is
little
control of
forest reserves, but the designation at
means
under
of
the legal instrument through which the
is
national parks
least
in
in
1973, and the Convention on Biological
international treaties are enforceable. All wildlife
that the areas are intended to remain
forest.
Conservation and research projects
There are many conservation projects

mostly coordinated
organizations.
work
by
International
the country, with
in
in
Nigeria,
nongovernmental
local
organizations
many
activities
close to the border with Cameroon.
the
Okwangwo
1998.'''
also
focused
AWWF program
The hunting
however This
of gorillas
is
increased conservation education
largely
in
is
now
due
Nigeria,
to
be-
Division of the Cross River
NP
in
until
Rural development, education, and pro-
tection efforts
adjacent
were carried out
Mbe Mountains, and
supported.
WWF
is
in
the park and
a gorilla
not currently active
but Fauna and Flora International (FFII
the gorilla's range, and
these was reported to have
of
killed eight gorillas.'"
much
are a current
increased on a road already
running through the Oban Division from Calabar to

the
Volker
in
the vicinity of
supported by the European Union was operating
females with young.
of
may be
concern. Traffic
in
NP
the northern parts of their Nigerian
range (especially Gashaka Gumti NP), but hunting
Infants are
are often not enforced. There
not generally taboo
southern Nigeria, although local taboos
Islamic areas
but
Oban
Conservation
habitat.
habitats.
the
farmland, plantations,
into
and bush-fallow.' The expansion
ape
Much
also occurs - by 1987, around 2k
ready been converted
palm
Nigeria,
in
are being actively logged.
all
illegal logging
percent
cover of
forest
2.6 percent.^ Tfiere are logging con-
although not
Plans for road development
Nigeria, with
in
of
census was
in this
is
area,
assisting
Sommer
Gashaka Gumti National

Park (Gashaka sector!
during the rainy season:
chimpanzee
habitat.
ginning with a Nigerian Conservation Foundation
(NCFI project that followed up on the 1989 survey,

followed by the
Global
Okwangwo program
Conservation
Organization
Nigerian
Conservation
recently,

is
an occasional report
hunters
in
but there
been
the
is
Okwangwo
Mbe
VWF-The
and,
most
Foundation-
(WCS) program. There
of a gorilla being killed by
Division of Cross River NP,
no direct evidence
killed at Afi or
of
in
of
any gorillas having
the last five years."
381
"
World Atlas
Cross River State Forestry Commission
the
develop
management
better
practices
Mountain. To the northeast, there
and behavioral research project
habituation
Gashaka Gumti NP,

program
jointly with the
Cross River
has a biodiversity
run
Nigeria,
Nigerian Conservation Foundation.
USAID and the US

its
WCS
southeastern
in
at
researchers from
involving
University College London.
research
Afi
research and conservation
gorilla
been proposed and are now
of
One would
Cross River
NP
stages of
at various
unite the
Okwangwo
south, would unite the
with Korup
NP
in
Oban
Division
Takamanda
with Cameroon's
NP
Reserve, one Gashaka Gumti
Cameroon's Tchabal-Mbabo NP, and
NP
southwest Nigeria and the Niger Delta,
In
surveys are urgently needed to assess the distribu-
abundance, and genetic
tion,
panzees. The
most
a third,
with
in
the
Cameroon.^'
be selected for a long-term research
The Pandrillus
more
effective,
force
them enhanced.
and the
Rehabilitation and
Drill
but
does
to
Breeding
orphaned chim-
them." Young
breed
not
life
at the
urban
facility in
km' enclosure
0.02
at
Calabar, then
There were 23 chimpanzees here

is
involved
in
for
move
Mountain, which
Afi
contributes to conservation education
center
in
in
the area.
2004. The
general conservation work
in
southeast Nigeria including creation of protected

areas,
community outreach, and education.
for all
Cross River
more
be
ef
priorities
for
(2003)'^
al.
latter
are the
action
for
are derived
chimpanzees and
gorillas.
recommendations from the
Second International Workshop and Conference on

the Conservation of the Cross River Gorillas held
Limbe Botanic Garden, Cameroon
in
in
gorillas,
major
three
the
first,
initiatives
establishment
Takamanda-Okwangwo complex,
for the
are
of
in
upgrading the protection status
ticular by
par-
of the
Takamanda Forest Reserve; second, the development of land-use plans for the wider TakamandaMone-Mbulu area in Cameroon, which would include a network of protected areas and corridors;
and
third,
plan for the conservation of the
Afi-Mbe-Dkwangwo area
of the
management
in
Nigeria, with a review
status for the
and the maintenance
of
Mbe Mountains
connections
forested
gorilla habitats.
been
has
all
recommended
that
remaining chimpanzee habitats
including, in particular, the
formal
Reserve);
Plateau
in
the
Nigeria,
in
community
Bayelsa state (the proposed
forests
Edumanom
remnant forests
of
the
in
Forest
Mambilla
Mbe Mountains
The Takamanda-Okwangwo
Taraba state; and the
in
Cross River state.
transboundary
2003.
Research
protected
would
area
benefit
gorillas.
Cross River gorilla management committees
Basic research into the ecology, distribution, and
should be established
population biology of the Cross River gorillas should
Capacity building for conservation and research
be expanded.
needed by the relevant
For
of
trade.
chimpanzees,
priority
actions
in
Gashaka-Mambilla area include supporting

conservation and research activities
Gumti NP, Nigeria.
It
distribution
in
the
existing
Gashaka
has been recommended that
basic surveys be conducted to assess
382
There should
awareness
raise public
protected area or reserve status be extended to
Sunderland-Groves and Dates 12003)" for
The
gorilla populations.
initiatives to
bushmeat
It
from Dates
and
conservation issues, including the impacts of the
between

The following
particular, protection
In
law enforcement measures should be strengthened
For
chimpanzees are quarantined and prepared
made
institutional capacity to en-
Cameroon-Nigeria transboundary protected area
Centre provides a sanctuary
to
effort.
Protection
""
forest
chim-
of
receive effective protection, and at least one should
recommended;
panzees,
affinities
populations should
viable
Protected areas and wildlife laws need to be
Three transboundary protected areas have
Wildlife
populations.
Fish and Wildlife Service fund
component.
discussion.
and potential connectivity between forests and
chimpanzee
is
to
at
chimpanzee
and numbers and evaluate existing
in
Cameroon and
institutions
in
Nigeria.
is
Nigeria and
Cameroon, including government departments,

universities,
and nongovernmental organizations.
Finally,
vital to
needs
it
is
into
the
incorporate local community
development
of
management
strategies, including the study of alternative liveli-
hood options.
AFRICA: Rwanda
FURTHER READING
Gonder, M.K., Oates,
chimpanzee
Ite.
J.F..
Adams, W.M.
U.E.,
A new West
African
state, Nigeria.
Applied
Disotell, T.R., Forstner, M.R., Morales. J.C, Melnick, D.J. [19971
subspectes'i'
Nature 388: 337.
119981 Forest conversion, conservation and forestry
in
Cross River
Geography ^8M: 301 -3U.

Oates,
McFarland,
J.F.,
history
GoriUa Biology:
Osemeobo,
K.L.,
and status
Is
Linder, J.M., Disotell, T.R. 120021 Tfie Cross River goriUa: natural
J.L., Bergl, R.A.,
neglected and
critically
120011
G.J.
Groves,
of a
traditional
endangered subspecies.
ecological know/ledge relevant
in
V.,
Adanu,
two years
J.,
Faucher,
I.,
J.L.,
Oates,
pp. 472-497.
131:
Nigeria?
in
203-210.
Fowler, A. 120041 Nigerian chimpanzees IPan troglodytes vellerosus] at Gashaka:
Primatologica 75
of habituation efforts. Folia
Sunderland-Groves,
Taylor, A.B., Goldsmith. M.L.. eds,
environmental conservation
InternationalJournal of Sustainable Development and World Ecology 8
Sommer,
In:
J.
151:
295-31
6.
120031 Protection strategies for Cross River gorillas. Gorilla Journal 27: 12-13.
http://www.berggorilla.de/engli5h/gjournal/texte/27crossrhtml.
MAP DATA SOURCES

Map
16.15 Great apes data are based on the following sources:
ButynskI, T.M. 120031 The chimpanzee Pan troglodytes: taxonomy, distribution, abundance, and conservation status.
Kormos,
R..
Boesch, C, Bakarr,
Butynski, T.M., eds,
M.I.,
In:
Wesf African Chimpanzees: Status Survey and
Conservation Action Plan. lUCN/SSC Primate Specialist Group. lUCN. Gland, Switzerland, pp. 5-12.
Oates,
J.,
Gadsby, L, Jenkins,
M.I.,
P.,
Gonder,
K.,
Bocian, C, Adeleke, A. 120031 Nigeria.
Butynski, T.M.. eds, West African Chimpanzees: Status Survey
In:
Kormos.
R..
Boesch, C, Bakarr,
Plan.
lUCN/SSC
Primate Specialist Group. lUCN. Gland, Switzerland, pp. 123-130.

With additional data by personal communication from Bergl,
Sunderland-Groves,
in
Takamanda
J.L.,
R. 120051
and from the following source;
Maisels, F, Ekinde, A. (20031 Surveys of the Cross River gorilla and chimpanzee populations
Forest Reserve, Cameroon.
In:
Comiskey,
J. A.,
Sunderland, TC.H., Sunderland-Groves,
Takamanda: The Biodiversity of an African Rainforest. Smithsonian
Institution.
J.L.,
eds,
ACKNOWLEDGMENTS
This country study draws extensively on the Nigeria chapter from the
Survey and Conservation Action Plan, Oates etal. 120021." and Sunderland-Groves and Oates 120031." Many thanks
Richard Bergl
ICity University of
(Zoological Society of
(Development
New
Yorkl.
San Diego), John
Nigeria! for their valuable
in
F.
Alexander Harcourt (University
of California, Davisl,
Oates (Hunter College. City University
comments on
New
of
Yorkl.
to
Bethan Morgan
and Jason
Sali
the draft of this section, and to Brigid Barry (Tropical Biology
Associationl for editorial assistance.
COMPILER
Monitoring Centre
REPUBLIC OF RWANDA
Nigel Varty

The Republic
26 338km*^
in
of
Rwanda
area.
It
is
is
rainy
mountainous country
located
in
the equatorial
highlands of the Western or Albertine
bordered by the Democratic Republic
Rift Valley,
of the
Congo
seasons
(lYlarch
Decemberl. The
Ivlay
to
official
English, and Kinyarviânda, and Christianity
majority religion.
Rwandan
been portrayed as consisting
of three
(84 percent), Tutsi (15 percent),
and Burundi.
and
has a temperate climate with two
historically the Tutsi
to
is
the
society has traditionally
IDRCl, Uganda, the United Republic of Tanzania,

It
and October
languages are Frencti,
and
groups: Hutu
Tvi/a (1
percent),
provided the governing
383
World Atlas
Map
in
Rwanda
tliis
country profile
1:
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in
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ra
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11
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oi
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Ci*
386
7-
>.,
v^
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,^\=^--
'
AFRICA: Rwanda
Rwanda gained Independence from Belgium

and was subsequently ruled by Hutu-
class.
in
1962,
dominated governments.'"
From 1972
party under President Juvenal
Hutu. Following the
fall of
ruled by one
Habyarlmana,
monarchy
in
the Tutsi
were both excluded from power
1961, the Tutsis
and subject
massacres. Many
to episodic
fled the
some gathered in Uganda to form an

(the Rwandan Patriotic Army,
country, and
armed movement
RPA)
Hutu dominance. The Rwandan
resist
to
Army
Patriotic
in
1990 but was
in
A power-sharing agreement
fought to a standstill.
was brokered
Rwanda
invaded
by countries
the region and signed
in
1994, but immediately after signing
Habyarimana was assassinated and
program
it
ranked 159th out
is
Hutus began. Hutu peasants,
incited to
kill
important sector, and

of the
in
177
of
the world."
in
by far Rwanda's most
is
2002 employed 90 percent
workforce and contributed 45 percent
About 42.6 percent
country
of the
permanent crops and arable
GDP.^
land.* Natural forests
cover about 462 km^ (1.8 percent! of land area,

according to FAG, although the area
forest suggests that this
km^
are a further 2 610
is
protected
of
an underestimate. There
of forest plantations, largely
eucalyptus and pine." Most of Rwanda's natural
of
forests
lie
in
the afromontane region
600
(1
by clearings (thought to be the result of
disturbance!
montane
human
and dense understory typical
forests.'
rate of forest loss
the region." Until recently,
in
there were four main
800 000 people were
[Nyungwe, Gishwati, Mukura, and Volcanoes!,
and over a million
refugees escaped from the country." The Rwandan
Army captured
Patriotic
ended
and
1994,
Government
earlier
of
the capital, Kigali,
genocide,
the
in
July
forming
the
National Unity to implement the
intervened militarily at this time. Fighting never-
1996 spilled
Rwandan
and around Rwanda, and
in
over
Patriotic
DRC
into
(then
Army backed
in
The
Zaire).
a rebellion there,
which destroyed the refugee camps and resulted
Mobutu
the overthrow of President
(see
DRC
country
profile!.
One
result
DRC
repatriation of refugees from
in
May 1997
was the mass
in late
Rwanda's western border

coalition with
DRC. The
Uganda,
led
Rwanda,
to intervene
fighting involved
DRC
again
initially
in
in
eastern
forces and their
Zimbabwean, Chadian, Angolan, and Namibian

on one side, and rebels supported by Uganda
and Rwanda on the other By the time that a
agreement was signed
in
autumn
the rebels controlled large areas
in
the north and
ceasefire
1999,
Further fighting and negotiations occurred,
east.
and a more complete accord was brokered by South

Africa
its
in
2002, allowing for
Rwandan withdrawal
of
troops from DRC, and disarmament, demobi-
lization,
In
lation
forests
the
of
Rwanda
in
Burundi
still
form one
of
all
Nyungwe
country.
NP
National Park [NP! and the adjacent KIbira
in
the largest remaining
blocks of lower montane forest
"
in Africa.'
and repatriation
of
Rwandan
growth rate
at 7.8 million in 2003, with a
of 1.8 percent.^ In 2002,
product (GDPI
was estimated
to
gross domestic
be US$1.7
gorilla
billion,
and gross national income (GNII per person was
{Gorilla
beringei benngeil
and the eastern chimpanzee {Pan troglodytes

schweinfurttiii] occur in
occur
Rwanda. Mountain
NP
the Volcanoes (Volcans)
in
Rwanda. The
gorillas
northern
in
comprise part
gorillas here
of the
larger single Virunga population, which includes
and
in
in
NP
the contiguous Mgahinga
the southern sector of Virunga
in
NP
Uganda
in
DRC.
This mobile population cannot be precisely divided
according
to
country,
has been censused
but
repeatedly since the 1970s [see Chapter
one
it
the
of
populations.
monitored
best
partial
by a
full
census
The census was

Gorilla
in
of
census
mountain gorillas was conducted
all
of
in
8!,
making
great ape
the
Virunga
2000, '^ followed
September and October 2003.
a joint effort of the International
Conservation
Programme
[IGCP!, Wildlife
Conservation Society WCS], Dian Fossey Gorilla
Fund International (DFGFIl and Europe [DFGFE! and

BerggoriUa &
Regenwald
population at 380, which
exiles.
the aftermath of these events, the popu-
was estimated

The mountain
those
1996.
By August 1998, continuing insecurity on
allies
west
the
in
montane
power-sharing agreement. France also
theless continued
in
located
of
The country has had the highest
propaganda, supported the extremists. At least

killed
to
4 500 m, typically 2 000 m!, and are characterized
full-scale
by ethnic
of
occupied by
is
President
genocide against Tutsis and moderate
of
Index
Subsistence agriculture
Rwanda was
to 199^,
US$230." The country's Human Development
only
(56
more
Direkthilfe.
was
It
put the
a 17 percent increase
gorillas! over the previous full
census
of
1989. Of these gorillas, about a third are estimated

to
be resident
in
Rwanda.'"
Eastern chimpanzees are restricted to the
southwest
of
Rwanda,
largely
in
Nyungwe NP,
385
World Atlas
although they formerly occurred in other forest

areas to the west' " Records also exist for a small
30-40 using the park some
area
consistent with an increased gorilla population
northwest,
the
in
Reserve area.' but

very few.* There
surveys, but
be
at least
in
if
Gishwati
the
in
Forest
they stiU exist here, they are
have been no recent national
1987-1989 there were estimated
to
numbers
it
the highest
any African nation,
of
is
human
population density
overwhelmingly agrarian,
and ranks among the 10 poorest nations
These factors
world.
result
the
in
acute land and
in
resource scarcity, and consequent pressure on

natural habitats.
Illegal
the country's main source of energy),

by honey gatherers),
all
mated
that
from
set
cannabis propagation, and
threaten Rwanda's forests.'
mining
fire [often
is
1990
50 km' or 3.9 percent of the total
year. This includes natural
"
It
is
esti-
an average
2000,
to
of
was cleared each
and planted cover, but
the figures of the Food and Agriculture Organization

of the United
Nations (FAO) indicate that there has
been an increase
in
Nyungwe
forest
cover over this
plantation
period, implying that the loss
is of
natural forests.'^
[now NP) covered
16 km'
forest covered 30
in
1996." There has been
the Volcanoes
was
lost
1996, and
1960,
but only
in
little
NPin recent years,^'
in
habitat loss
in
but half the park
during the late 1960s to early 1970s.^'^
2004, there
In
km'
UO km'
km' by
1960, but had been reduced to 945
Mukura
was
proposal to route electricity
pylons through the park to the radio antennae on

the
summit
In
to
be
of Karisimbi
influx of
Rwandan
Rwanda
very small and at risk.

to
have declined
this
the
in
is
more
a reflection of a
There does seem
field.
have been a marked reduction
in
with chimpanzees lost from the center-west region

of
Rwanda
since 1940, and no records since 1983
Although
the
bushmeat trade
following the genocide
generally
are
gorillas
hunted for food
not
Rwanda. Two nursing female
gorillas
were
Rwanda on May
in
sell the infant."
baby
is
to
no established trade
in
however, and a possible threat from
gorillas,
poaching
is
NP
poachers hoping
9 2002 by
There
in
killed
and a young male abducted from the Volcanoes
inferred only from the fact that
market
prices are quite high." That few young gorillas have
been taken
in
recent years, even during periods
of conflict, reflects the dedication of
zations
much
in
maintaining patrols
greater threat
ungulates by people
NP
Volcanoes
NP
and international nongovernmental organi-
staff
in
is
of
key gorilla areas.
the setting of wire snares for
living
adjacent to the Volcanoes
Rwanda, and by professional poachers. Two
veterinary surgeons are permanently employed at

the park to monitor the health of habituated groups
of
mountain
gorillas,
and
There
is
high
remove snares from the

"
to
human
population density
around Nyungwe" and snares are common, with

several thousand
being
guards from the Office
collected
of
numbers
There
is
of
annually
by
Tourism and National
Parks [ORTPN). Those that remain
Virunga
increased
1994," chimpanzees and
in
undoubtedly pose a threat
of
in
several other areas.'
deforestation of large
National Park.
to
range, however,
refugees led to
areas
in
over the last 50 years, though
whether
not clear
Is
were thought
the Volcanoes NP, with an additional

El(zabeth A. Williamson
The
still
limbs of injured animals."
Volcano."
1981, 110 mountain gorillas
living in
in
lack of observers
clearing for subsistence
crops, tree felling for timber and fuel [firewood
is
in
encouraging, the global mountain
is
gorilla population is
Chimpanzees are thought
THREATS
time." By 2001,
animals.' This
the Virungas as a whole over the last 20 years.''"
While this
500 chimpanzees remaining.''
Rwanda has
of the
was 129
the estimated population
to
in
the forest
chimpanzees, but the
deaths and injuries are unknown.'"
no sign that Nyungwe chimpanzees have
been captured
for the pet trade.'" Mining
harvesting are ongoing
in this
and forest
park, and forest fires
are frequent.''
Mountain gorillas can suffer from many

the
same diseases
tourist
groups, researchers, and local people
believed to have caused a

illness in the Virungas,
those of hunting."
386
of
as humans." Close contact with
In
number
of
outbreaks
which have impacts
is
of
rivaling
1988. for example, measles or
AFRICA: Rwanda
a related morbiUivirus killed six liabituated female
two."
l<illing
"
gorillas. ''
mountain
affected 26 of
1990. bronctiopneumonia
In
35 gorillas
tfie
When
a tourist group,
in
the security situation allows, 70
percent of the Virunga gorillas are visited daily by
more than 70
guides,
and a similar number
tourists
continued daily exposure
to
people and their diseases
is
major potential
of
rangers, and researchers.'' This
porters,
Groups
threat.'
numbers
large
considered
of
be a
to
chimpanzees
of
habituated for tourism and research are similarly

vulnerable' and those
be at
Nyungwe NP
the
in
could
risk.
The war and genocide

massive increase
Rwanda caused
in
human
in
A juvenile mountain
across the
traffic
Virungas. followed by a sustained military presence.
known
Parasites
sequently found for the
time
first
mountain
in
Between 1992 and 2000, 12-17
gorillas."
died
humans were sub-
infect
to
gorillas
from military gunfire or explosions
the
in
Virungas,'" representing between 4 and 5 percent

of the
1989 population. The recent conflicts and
subsequent resettlement
sands
ape
for instance,
this
tens of thou-
refugees and internally displaced people
of
have also resulted

of great
many
of
the
in
habitat.
and degradation
The Gishwati Forest Reserve,
was almost
and only
period
loss
forests.
degraded fragments
which
(DoFl.
ratified or
Biological
acceded
Diversity
International Trade
in
to the
1996;
the
in
1980; the
Convention on
UN
Forestry
of
Environment, Forestry, Water, and
of
Lands,
Natural Re-
sources (MINITEREl. On the whole, the depart-
ments deal with
policy development,
and the
agencies with implementation.''
The Department
responsibility for
of
conservation.
reserves
of forest
Department
Environment has overall
biodiversity
The
of Forestry.
The
the responsibility
is
Office of
Tourism
and National Parks, however, based within the
Convention on
Endangered Species
in
Fauna and Flora ICITESl
Department
alongside the Department of
sits
Environment (DoEl within the Ministry
of the
LEGISLATION
Prime
02/88 and
Decision
organization of the
the
management
remain.'
Rwanda
Ministerial
Ministerial Order 72/03 of October 14 2002 detail
encroached during
totally
tiny
gorilla.
Charles Mayhew/Tusk Trust
of
Wild
Convention
Ministry of
tion,
Commerce,
management
for
Industry, Investment
Promo-
Tourism, and Cooperatives, has responsibility

of
parks and matters
national
relating to ecotourism.
The Volcanoes and Nyungwe
the African
NPs
of
Nature and
Tourism and National Parks, whereas the Mukura
Natural Resources
in
Convention on
and Gishwati Forest Reserves are the responsibility
Migratory Species
2005; and the World Heritage
to
Combat
Convention
Desertification
in
in
1998;
in
1979;
the
2000. The country also participates
in
UNESCO's Man and Biosphere (MABI Programme,

underwhich
a 151
km' Biosphere Reserve has been
of
are therefore the responsibility of the Office of
Department
the
of
Forestry.
The
Rwanda
Environmental Management Authority IREMAI has

also
been established, and
charge
of a
wide range
of
expected
is
to
take
environmental projects.*
designated at the Volcanoes NP.
Chimpanzees and
law,
and
restricted.'^
national
gorillas are
and
protected by
international
trade
is
Ordinance 18/6/73, modified by Law-
Protected areas
There are four main categories

in
Rwandan
of protected
area
law: national parks, hunting reserves,
Decree 26/4/1 973 and Law 34/2000, established the
special reserves, and forest reserves. There are
Tourism and National Parks, and governs
currently three national parks, including Volcanoes
Office of
the creation and functioning of protected areas
NP and
and hunting arrangements. Forestry matters are
three forest reserves, but no hunting reserves or
governed by Law 47/1988, which sets out the basic
special reserves.
legislation
concerning conservation and use
of
of the
the recently designated
country
was
'
Nyungwe NP, and
Until 1997. about 15 percent
included
in
national parks and
387
World Atlas
NP
reserves, but the Akagera
reduced
about 40 percent
to
subsequently
2003, bookings for gorilla tourism were at about
of its original size to
60 percent capacity," while visitors to Volcanoes
Vi(as
allow resettlement of refugees. Today, the protected
area system covers about 8 percent of the country.
The Virunga NP on the borders
DRC was
day Rwanda, Uganda, and
mountain
for the protection of the
the
national park
first
was established as
in
of
929.
created
gorilla,
in
1925
and was
NP
the separate Volcanoes
Rift
International
reduce pressures on
of
Nyungwe
to
be widespread across the
reserve, especially
is
the western section, and the
in
population,
at
living
one
of
Some
in
the 980
km'
most recent estimate dates from the
more than
000 were thought
to
be present
Chimpanzees apparently
the early 1980s."

in
highest
the
recorded for this species.^'
100-200 chimpanzees may remain

park, but the
survive
(now NPI
forest
thought to support a relatively large and
altitudes yet
In
typically
the nearby 6
still
km' Cyamudongo Block
the southwest of Nyungwe, which has
Incorporated into the park.'"
common
among them
Chief
is
strategic frame-
and a number
NP
the Volcanoes
IGCP, a joint
of
area.
of
initiative
the African Wildlife Foundation (AWF). Fauna and

International
Flora
and
IFFII,
WWF-The
Global
to
now been
The park has been
ing
Rwanda
in
since
1978.
IGCP continued
to
provide technical and material support to the park
throughout the conflict
emergency funding
the
in
basic
for
salaries and rations tor the patrols,
medical supplies, rehabilitation
and clearing
of
Including
1990s,
park operations,
equipment and
of infrastructure,
mines. Other groups Involved
with gorilla conservation
in
Rwanda
include the
Dian Fossey Gorilla Fund International and the

DIan Fossey Gorilla Fund Europe, which support
research and community awareness, and work
in
Volcanoes NP, the Karisoke Research Center, and

the
Mountain Gorilla Veterinary Center (MGVCl.
the past by mining (particularly
Together these provide technical and financial
for gold), road building, selective logging, reforest-
assistance worth around US$1 million per year to
heavily disturbed
ation with
In
Chimpanzees were also
^'
known from
viously
1985," but after the
km'
in
the
Office
of
Tourism and National Parks
the
park,
around
Its
WCS
and
borders.
development
local
for
'"
has a long-standing Project
Conservation of
Nyungwe
about US$200 000 per year
Conservation action
and National Parks
Wildlife tourism, particularly visits to the Virunga
seen as a key tool
is
conservation areas
NP
Volcanoes
million
Tourism
1989,
In
at
in
direct
Nyungwe
protecting forest
Rwanda. Tourism
provided
in
in
Rwanda
with
In
the
US$4-6
and indirect revenues.
provided around US$0. 5-1.0
During the
of
for
the
Forest (PCFN), which
to the Office of
Tourism
management, research,
for
tourism development, rural public awareness, and

staff training at
far
Nyungwe
NP. Ninety guards have so
been trained under the project." The project
established a research station and tourist

at
Uwinka
forest
in
in
the northwest sector of
the 1980s. These
facilities
Nyungwe
were looted during the
genocide, but were saved from destruction by the
million in 1990.
Volcanoes
at
activities
provides technical and financial assistance worth
survives here.
gorillas,
for
management, research, and veterinary support
of this
doubtful that the species
is
it
13 remained
at least
conflict only 6
civil
remains and
pre-
the GIshwati Forest Reserve
where
(near Ruhengeril,
forest
and encroachment by
exotic species,
farmers."'
local
conflict, the
NP was
headquarters
ransacked and destroyed.
the staff were
evacuated. Tourism
killed
Is
of
the
Some
and others temporarily
now
security concerns remain
388
Tourism and National
Office of
works with nongovernmental orga-
Conservation Organization, which has been operat-
A 1999 survey
980s;''
The Rwandan
Parks
In
to try to
is
local
In
development projects."
such organizations work
found chimpanzees
spent on protecting the park and investing
goals,
'on-farm resources'
tourist per
contact time with gorillas.^' The income
nizations to establish a
the park.
viable
US$350 per
to
work and conservation
IIGCPl are working with local farmers to develop
reserve
of
2004
in
60 km'. Of the hundreds
is
species found here,
Western
hour
armed guards. Fees
escorted by heavily
many are unique to the

region. CARE International and the
Gorilla Conservation Programme
current area
Its
of the present-
NP were
were increased
The Rwanda portion
Africa.
in
'"
Increasing again, but

a
major deterrent.
In
French military Intervention

since
In
Rwanda.^" They have
been repaired and small streams
searchers and tourists have begun

Following
the deforestation
of a large area In the
of
re-
to return.
and
MIkeno sector
of
cultivation
Virunga
NP
AFRICA: Rwanda
DRC In 2004, the United Nations Environment

Programme lUNEPI, WWF, IGCP, the European
in
Union, and Frankfurt Zoological Society released
emergency funds
support the Congolese Institute
to
Nature Conservation (ICCNl
for
the
w^lth
con-
struction of a dry stone wall to help restore the

integrity of the parl<
km
boundary." By August 2004, over
had been completed, and the work
of the wall
continues with a workforce of more than 2 000
Rwandan
people. Six
the wall, which
of
associations are building part
Guards
runs along the international
boundary. The construction of the wall clearly
demarcates the boundary

expected that
as the
it
will,
the park and
of
come
over time,
to
it
new
is
be accepted
'^'
Ian
vation of protected areas.
The measures proposed
include development and Implementation of land
base for most research within Volcanoes
use and management plans for each protected

area, continuing research at protected areas inclu-
the late 198Qs due to political and
ding Inventory studies, and the sustainable use of
established
site until
In
she was murdered there
murdered, resulting
unrest.
civil
It
1967 and DIan Fossey ran the
three Spanish medical workers

in
in
1985. In 1997,
in
Ruhengeri were
the departure of expatriate
many
Great insecurity followed and
staff.
uniforms.
Redmond/UNESCO
NP, although research has been intermittent since
was
Volcanoes
Action Plan INBSAP) calls for Improved conser-
limit of cultivation.
Much research has been conducted on the

^^" Karisoke is
Virunga mountain gorillas.'^- '
the
at
National Park receive
Including several Karisoke staff,
people.
were arrested and
development
biodiversity including
In
November
2004,
of ecotourlsm.'
Rwanda became
the
of
first
the 23 great ape range states to officially endorse
and distribute
plan.
The
chief
its
great ape
national
recommendations
survival
of this plan
are
outlined here.
imprisoned.^"
Organizations Involved
in
2003 census
the
park authorities (Rwanda's Office

National Parks, the
Uganda
and the DRC's ICCNl, the
of
Tourism and
protect and conserve Volcanoes NP, at least
Authority,
within the current limits of the park, by dimi-
W/lldlife
for
Institute
Tropical
Forest Conservation (ITFCI of Uganda's Mbarara
Dian
the
University,
Fossey Gorilla Fund, the
Mountain Gorilla Veterinary Center, WCS, and the
Max Planck
Gorilla
Institute for Evolutionary Anthropology.
and habitat monitoring teams have been
established
in all
the Virunga parks by
IGCP and the
There are no great ape sanctuaries
in
Rwanda,
but the Mountain Gorilla Veterinary Center, estabin
1986 near the Volcanoes NP, looks after
the health of wild gorillas and helped to care for
baby gorillas confiscated by the authorities
One
Infant
was reintroduced
one was reared
nishing poaching and other illegal activities;
increase the benefits to local communities
from conservation, and increase local support

for conservation;
increase awareness of conservation Issues at

the local and political levels;
continue and Increase research and monitoring for the gorillas
park authorities."'
lished
For gorillas:
Virunga gorillas included IGCP, the three
of the
In captivity,
to its
in 2002.''
natal group, and
improve human,
and
their habitat;
gorilla,
and livestock health
standards; diminish disease

illas
through control
livestock Interaction;
of
among
the gor-
zoonoses and
gorilla/
and
Increase tourism revenue through sustainable
ecotourlsm.
but both died.
For chimpanzees:

A workshop
of
Including a
In
held
in
July 2003 led to the production
national great ape

list
of potential
survival
plan
(NGASPl,
conservation projects.''
addition, the National Biodiversity Strategy
and
reduce the area
of
ensure no further loss
degraded habitat and

of habitat;
promote, implement, and enforce laws and

cies regarding
chimpanzees and
poli-
their habitat;
389
World Atlas
increase awareness
importance
and pride
of,
in,
covering the whole range of the Virunga gorillas."
the
both chimpanzees and their
of
Further investigation into the possible effects of
and transmission
stress
improve community livelihoods around chim-
human
panzee habitats through conservation;
recommended'' " and recommendations and
create
greater understanding of status,
improve
developing
is
framework
regional
is
NP and
Ruhengen
the
apes has been
of the
mountain
improved conservation awareness and
revenue-sharing programs, and better information
for
on the numbers and distribution
a proposal
of
chimpanzees
Nyungwe NP have also been recommended.

A census of Nyungwe chimpanzees, funded by
develop a tourism development plan for the
Volcanoes
to
Continued close monitoring

gorillas,
conserving gorilla habitat, and there
suggested.""
of
Nyungwe and Cyamudongo.

IGCP
of
habituated apes has been
humans
transmission from
management
and
protection
contact with
best practice' code to reduce the risk of disease
trends, and biology of chimpanzees; and
to
disease from close
habitat;
in
region. In addition,
the park authorities of the three countries involved
WC5
have accepted the creation
early 200A.
of a transfrontier park,
and the University
of Antioch,
was ongoing
in
FURTHER READING
DoE
and Action Plan
120031 National Strategy
Environment, Ministry
of
for the Conservation of Biodiversity in
Lands, Resettlement, and Environment,
Rwanda. Departnnent
of the
Rwanda. http://www.biodivorg/doc/
Kigali,
world/rw/rw-nbsap-01-en.pdf. Accessed March 31 2003.
Kanyamibwa,
S. 119981
Impact
of
war on conservation: Rwandan environment and
wildlife in agony. Biodiversity
and
Conservation!: 1399-U06.
Montort, A. 11992) Premiere
Robbins, M.M., Sicotte,
P.,
liste
commentee des mammiferes du Rwanda. Journal
of African Zoology '[Qb. 141-151.
Stewart, K.J., eds (2001) Mountain Gorillas: Tfiree Decades of Research at Karisol<e.
MAP DATA SOURCES

Map
16.16 Great apes data are based on the following source:
Butynski, T.M. I2G01) Africa's great apes.
Stevens,
E.F.,
Arluke,
A.,
In:
eds. Great Apes
and Humans: The
T.S.,
T.L.,
Norton,
B.,
Rowan,
A.,

With additional data by personal communication from Plumptre,
Kalpers,
in
J.,
Williamson,
E.A.,
A.,
A. [200'1)
Nzamurambaho,
A., Lola, N., Mugiri, G. 12003) Gorillas
the crossfire: population dynamics of the Virunga mountain gorillas over the past three decades. Oryx 37: 326-337.
Lee, P.O., Thornback,
J.,
Bennett, E.L. 11988) Threatened Primates of Africa. The
lUCN Red Data Book. lUCN,
Gland,
Switzerland and Cambridge, UK.
MINITERE
12004)
Rwanda's National Great Apes Survival Plan 200i-2009.
Forestry, Water,
Plumptre,
A.J.,
and Natural Resources, Republic
Masozera, M., Fashing,
P.J.,
of
McNeilage,
surveys of the Nyungwe Forest Reserve
in
Final draft. Ministry of Lands, Environment,
Rwanda.
A.,
Ewango, C, Kaplin,
S.W. Rwanda.
WCS
B.A., Liengola,
Working Paper
19.
I.
12002) Biodiversity
http://www.wcs.org/rnedia/
file/workingpaperl 9.pdf.
ACKNOWLEDGMENTS
Many thanks
to
Alexander Harcourt lUniversity
of
California,
Davis),
Annette Lanjouw llnternational Gorilla
Conservation Programme), Andrew Plumptre IWildlife Conservation Society), Elizabeth A. Williamson

Stirling],
and an anonymous reviewer
for their valuable
comments on
AUTHOR
Nigel Varty,
390
UNEP World
lUnive.'-sity of
AFRICA: Senegal
REPUBLIC OF SENEGAL
Edmund McManus

The Republic of Senegal
of
West
Africa, bordering the Atlantic
Guinea-Bissau and Guinea
penetrates the central axis
with Senegal (under the
Senegal
to 1989.^
1982
from France
to the east.
in
1960,
The Gambia
brief political union
name Senegambial from
itself
its
Mauritania
the country along the
of
and formed a
River,
Ocean, with
to the south,
and Mali inland
to the north,
Gambia
grated to urban areas, such as the capital Dakar,
the westernmost country
is
became independent
capital
Dakar having
until
been the administrative headquarters
then
French West
partly
Africa.'
A succession
of
moderate,
of
democratic governments held power during
the 1960s and 1970s, and moderate, left-leaning
governments won
series of regular elections
through the 1980s and 1990s, with the opposition
then winning
2000 presidential and
the
2001
legislative
elections.'
There has been an armed
separatist
movement
in
Casamance
the
region of
southern Senegal since 1982. This region

richest agricultural area
of conflict.
It
is
the
Senegal, but as a result
in
lacks basic infrastructure and good
The
area of Senegal
total land
rainfall
is
ranges from 200
mm
in
temperature range
of
northern regions to
600
search
of
employment, agriculture remains the
millet,
The main crops are maize,
mean
daily
a dry
season from November
rainy season from |vlay/June to
196 190
mm
in
km^
the
nomy has
benefited from a stable microeconomic
policy since the late 1990s, but the country's gross
domestic product [GDP]
in
2002 was US$4.9
unemployment, high
social indicators (high
some poor
acy rates, and

cially
rural areas].
in
was
2001, Senegal
In
illiter-
health statistics, espere-
classified as a least developed country (LDC] by the
UN and in 2004 was ranked 157th

Human Development Index
(out of 177) in
the
Programme
Nations Development
of
United
the
(UNDP).''

The only great ape
be recorded
to
Senegal
in
is
the western chimpanzee [Pan troglodytes verus].

is
It
restricted
The
country.
the
to
distribution
was
carried
stage
April/May and a
region
part
of
of
the
systematic national survey of
first
chimpanzee
administrative
the southeastern
in
18-35C. There
October" There are
billion,
US$470 per person." Senegal has

many structural economic problems and poor
or less than
2003-2004 and the results were
is
The eco-
chief export crops.
and peanuts are the
the south, with a
to
rice,
sugarcane, alongside livestock, while cotton
Tambacounda,
links to the rest of Senegal.
Annual
in
chief industry.""
at the
out
in
analysis
2004.'
in
data
Existing
suggest that approximately
percent of the country's chimpanzees are
10
is
thought to live in the fJiokolo-Koba National Park

(NP),' " with the remainder of the population being
the Sahelian zone, with vegetation dominated by
distributed to the south and southeast of the park
Acacia spp. and annual grasses. The central zone
(Map
three main zones of natural vegetation, correlated
with rainfall, from north to south. Northernmost
is
Sudanian, with a range
wooded savannas
of vegetation
to dry forests.
types from
Southernmost
is
the Guinean zone, characterized by less dry forests,
with
gallery
forests
along
the
river
courses.
Senegal replants forest trees on about 300 km^

land each year, though
many young
of
trees die due to
in
16.17). Of 10
areas around the park surveyed
2000-2001, four were found
to
have relatively high
concentrations of chimpanzee nests: Bandafassi,
and Tomboronkoto.'^ The
Segou,
Fongolembi,
Diarha River area serves as an important dry-
season refuge
All
for several families of
chimpanzees.
populations are small.
lack of follow-up care.'

In
mid-200i, there were an estimated 10.8
THREATS
Senegal's chimpanzee
million people in Senegal, with a population growth
Although
rate of 2.6 percent.^ There are several ethnic groups,
population has declined historically,' recent trends
including Wolof (43.3 percent], Pular 123.8 percent),
are unknown.
Serer IU.7 percent], Diola

13
13.7 percent],
Mandinka
percent], Soninke 11.1 percent], and others (5.4
percent], but the population
is
95 percent Muslim.'
Even though many young people have mi-
mum
of
countries
it
clear that
is
In
300,'
made
1979
but
it
was estimated
at a
maxi-
national estimates for other
at this
time have since proved
to
be underestimates. The most recent population

estimate, published
in
West African Ctiimpanzees:
391
World Atlas
Map
r)
a>
id
Ol
<)>
(U
n n n
(1)
!>
.>
r*
<u
0)
o
^
1^
a.
u.
u.
t
o
^
F
f>
^ ^ ^
^
t:
Q.
(0
392
in
tiJ
'^y
in
Senegal
Data sources are provided at the end of this country profile
AFRICA: Senegal
Status Survey and Conservation Action Plan,
there
200-400 chimpanzees
are
range
country.' This
derived
is
living
is
that
Natural Resources. The country provides chim-
in
the
panzees with complete protection under the Code
by extrapolating
Hunting and the Protection
for
Fauna.' Although
of
density figures taken from specific field sites within
it
Senegal, estimates derived
capture them for approved scientific purposes, no
other countries with
in
theoretically possible to obtain a license to
is
similar habitat types, and responses to question-
such licenses have been issued. Forest protection
naires and Interviews.' "
provided for under the Forest Code. National plans
" Widespread habitat
'^
destruction, fragmentation of remaining forest, and
food sources are
all
to
National
d'Amenagement du
threats to Senegal's chimpan-
National
zees, however, so the apparent population stability
cannot be assumed
be
to
be the most significant
most
clear that
of
of
these threats.
chimpanzees
area
outside the protected
live
network. They typically inhabit small areas of forest
under increasing pressure, including the
that are
them
destruction of forest corridors that connect
other forest blocks.'
is
It
Le Plan
Terntoire,
I'Environnement;
La
Strategie Nationale et le Plan d Action pour la

Biodiversite;
la
Le Plan
and
d'Action National de Lutte contre la Desertification.
2003,
In
is
It
small population
of the country's
pour
d'Action
Conservation de
real.
Habitat degradation and alteration are thought

to
environment include: Le Plan
the
relevant
over critical water and
humans
competition with
Is
moratorium was placed on the
granting of quarrying permits

in
an attempt
persuade companies involved

to
move
in
forest reserves,
reduce deforestation. Efforts
to
in
to
existing operations
out were also underway.'
to
estimated that there are
Protected areas
62 050 km^ of forest remaining, but most of this
Senegal's national parks are
managed
unsuitable for chimpanzees.^ The deforest-
Direction des Pares Natlonaux.
The NIokolo-Koba
area
Is
ation rate
about 450 km'
Is
(0.7 percent!
Chimpanzees and humans compete

honey and the
Including
wild foods.
per yean'
NP, which
for various
Heritage
of
fruit
the
climber Saba senegalensis lApocynaceael.'^ As the

dry season progresses,
dry up and
also
many
natural water sources
some areas chimpanzees and humans
in
compete
for
access
to
those tew that remain.
Islam, the majority religion

hibits
consumption
of
Senegal, pro-
In
chimpanzee meat, and
tional beliefs also forbid the killing of
on the grounds
of their
unique position
tradi-
the
chimpanzees
in
Senegal
at present,
Guinea, where hunting

Since
Project
captive
1997,
the
Is
more
although the
baby chimpanzees
Investigation of the
In
of
Niokolo-
the border
five
reports of
Senegal.
cause and scope
Further
of the
problem
Senegal
to
moves
to treat
Is
contain
NP
contiguous with the Badlar
these two
Biosphere Reserves as a Niokolo-Badiar trans-
authorities already cooperate to

scientific
some
in
in
the two parks appear to be isolated
Niokolo-Koba
NF was
created as a hunting
1951, a faunal
reserve
in
1926, a forest reserve
reserve
in
1953, and a national park
was enlarged by
In
in
1954, and
a succession of decrees In 1962,
1965, 1968, and 1969.

level to the
Its
extent
and technical matters." The chimpanzee
summit
of
It
rises from just above sea
Mount
Asslrik at 311 m, and
ecosystems comprise about 55 percent grass-
land,
37 percent woodland,
and 3 percent
forest.
percent bamboo,
The vegetation varies from
southern Sudanian type
has been recommended.'
km' and
9 130
of
In
boundary protected area. The two management
populations
Rehabilitation
The Gambia has received
In
Is
from one another.'
prevalent.
Chimpanzee
It
among
other
area
Guinea, and there are
chimpanzees
home ranges of populations to the south

Koba NP are thought to extend across
Into
has an area
only protected
primates." Hunting does not appear to be a threat

to
also a Biosphere Reserve and World
is
Site,
chimpanzees.'
In
by the
to
Guinean, with savanna
predominant, more luxuriant vegetation along the
LEGISLATION
Legislation
Senegal
is
national Trade
party to the Convention

In
Endangered Species
of
on InterWild Fauna
course
of the rivers,
and a varying cover
of trees
topography and
and
The
bushes according
to local
only people to
within the park are forest guards
and workers
live
at tourist
soils.
camps.
and Flora ICITESl, the Convention on Biological

Diversity,
the World
the Convention
on Migratory Species,
Heritage Convention, and the African
of
Nature and
Conservation and field projects
The Programme d'Education
et
de Recensement
des Chlmpanzes du Senegal IPERCSI has carried
393
World Atlas
over resources, especially Saba senegaiensis!" The
Diarha River
groups
the district of Salemata has several
in
chimpanzees and, following the PERCS
of
surveys, has been selected as a second site for
long-term monitoring
to
determine the number
chimpanzees using the area and
of
to identify their
migration routes.'
There are no known sanctuaries, rehabilitation

centers, or reintroductlon sites for
in
Senegal. The zoo
chimpanzees
Dakar has a small captive
in
collection.
Officials at
an
illegal
logging operation.
The following recommendations were made
chimpanzee surveys
out
understanding
their
obtain
to
of their distribution,
complete
an estimate
of
Action
numbers, and more information about threats

Key populations were moni-
to their populations.
tored over a year to identify migration patterns and

sites.
Education activities were
in critical
areas, and a second phase
dry-season refuge
also carried out
is
proposed
concentrate on educational activities
to
and solutions
and competition over
conflicts
to
PERC5 works
Plan.''
Senegal,
sidered priority sites
protection
of
The Niokolo-Koba NP
with the
first
their
for
of
is
the site of the only
chimpanzee ecology
in
Senegal,
McGrew
data having been collected by
and colleagues
in
the 1970s."
established a research site
'*
In
May
2001, Pruetz
the Tomboronkoto
in
amended
chimpanzees
conservation,
purposes as
long-term study
of special attention.
chimpanzee populations migrating
should be
US nongovernmental
need
across the Senegal-Guinea border The relevant
of
Animals, a
in
Senegal should collaborate with Guinea on the
legislation
of
for
to prohibit the
capture
any reason other than
thus excluding
scientific
a justification. Sustainable solutions to
competition between
water and the
humans and chimpanzees
fruits of
sought. More education
work needs
to
be carried out
throughout Senegal, emphasizing the role
of habitat
destruction as a threat to chimpanzee survival.
ecotourism may be
Chimpanzee habituation
chimpanzees, following up on work carried out
detrimental to the already fragmented groups
McGrew,
"
Assirik.'^'
'
colleagues
at
Mount
The Tomboronkoto population
humans and
close to
and
Tutin,
they are
sometimes
lives
in conflict
over
Saba senegaiensis should be
region to study the ecology and behavior of savanna'
by
In
or that could
support chimpanzee populations should be con-
and trade
organization.
contain
locations that
all
alongside the Direction des
from Friends
H'est
With only 200-400 chimpanzees remaining
Pares Nationaux, and receives financial support
water^
in
and Conservation
for
In
the
country, so a multidlsciplinary board of scientists and
conservationists should consider the Issue on a case-
by-case basis before such
initiatives
are approved.
FURTHER READING
l^cGrew, W.C, Baldwin,
Africa.
Journal of
P.J., Tutin,
Human
C.E.G. 119801
Pruetz, J.D., Merchant, L.M., Arno,

in
Chimpanzees
In
a hot, dry and
open
habitat:
Mt
Assirik. Senegal,
West
Evolution 10: 227-244.

J.,
McGrew, W.C. (2002) Survey
of
savanna chimpanzees IPan troglodytes verus]
Senegal. American Journal of PrimatologySS: 35-43.
UNESCO
120021 Biosphere Resen/e Information. Senegal: Niokolo-Kcba. http://viww2.unesco.org/mab/br/brdlr/
directory/biores.asp?mode=all&code=SEN+03. Accessed November 17 2004.
MAP DATA SOURCES

Map
16.17
Butynski, T.M. (20031 The chimpanzee

In:
Kormos.
R.,
Pan
Boesch, C, Bakarr,
following sources:
troglodytes, taxonomy, distribution, abundance, and conservation status.

M.I.,
Butynski, T.M., eds. West African Chimpanzees: Status Survey
396
and
AFRICA: Sierra Leone
Carter,
Ndiaye,
J..
T.M..
S..
Pruetz,
McGrew, W.C. 120031 Senegal.
J..
In:
Kormos,
Boesch. C. Bakarr,
R.,
Butynski.
M.I.,
West African Chimpanzees: Status Survey and Conservation Action Plan. lUCN/SSC Primate
eds,
Specialist Group. lUCN, Gland. Switzerland, pp. 31-39.
ACKNOWLEDGMENTS
This country study draws extensively on the Senegal chapter from the
Survey and Conservation Action Plan. Many thanks
comments on
to
lUCN/SSC West African Cfiimpanzees: Status
Georges Grepin and an anonymous reviewer
for their valuable
COMPILER
Edmund Mcfânus, UNEP World Conservation
Monitoring Centre
REPUBLIC OF SIERRA LEONE

Edmund McIvIanus

The Republic
Leone
of Sierra
Guinea and Liberia and

countries
71 620
West
in
km'
of the
one
is
between
situated
is
the smallest
of
Africa, with a land area of
divided into the Eastern, Northern,
Southern Provinces and the Western Area.
wooded
about
hill
in
seasonal tropical climate, with
May
December and
to
income.
to April,
when
dry dust-laden harmattan winds blow
from the Sahara. Mean annual
in
from
830
mm
in
places
in
range
is
West
Africa,
UK
it
ranges
savannas
one
of the
to
wettest
and the annual temperature
Leone became inde-
21-36C.' Sierra
pendent from the
rainfall
northern
the
mm on the coast, making
5 230
season from
a rainy
season from December
a dry
The population was about
5.9 million in 2004,

In
2002, gross
domestic product (GDPl was US$782.9
million, with
with a growth rate of 2.3 percent.'
income (GNU
a gross national
per person.'^ English

(although
limited
to
is
the
of less
the
literate
English-based Krio ICreolel
is
than
official
US$UO
language
minority),
but
understood by 95
percent of the population. The main ethnic groups
Temne (who
are
Mende (who
are dominant
are dominant
in
in
the north] and
the south], at 30 per-
cent of the population each, with Islam as the main

religion (60 percent of population); sizeable
ities
or
a
minor-
practice traditional beliefs (about 30 percent)
some form
major
of
role in the
Christianity.'^
economy
Agriculture plays
with about two thirds
Diamonds,
of Sierra
Much
trade
armed
sustain
and
fish
sub-
in
(titanium
rutile
have con-
Leone's exports historically,
1991-2002
in
of
more than
civil
war
2 million people (well
over one third of the population],
now refugees
of a
UN
many
whom
of
are
neighboring countries. With the
in
peacekeeping force and contribu-
tions from the international

bilization
to
tens of thousands of deaths and the
in
displacement
support
in
war between the government and
civil
the Revolutionary United Front (RUF]. The

resulted
of
diamonds was used
conflict'
factions and their mercenaries
community, the demo-
and disarmament
United Front and
1961.
in
diamonds have become the major source
but
has
the west. Sierra Leone has a
most
tributed
country, upland plateaus, mountains
mangrove swamps
farming.'
oxide], bauxite, cocoa, coffee,
and
It
the east, and a coastal belt mostly covered by
in
working-age population engaging
sistence
Civil
of
the
Revolutionary
Defense Force combatants
have been completed. National elections were held

in
the government
May 2002 and
slowly to re-establish
withdrawal
in
of
most
Sierra Leone
in
its
of the
is
continuing
authority The gradual
UN
peacekeeping mission
2004 and the security situation
neighboring Liberia
may
in
present challenges to the
continuation of Sierra Leone's
Fighting
stability.
among
disparate rebel groups, warlords, and youth
gangs
in
Guinea, Liberia, and Sierra Leone has
created insurgences, street violence, looting,

trafficking, ethnic conflicts,
areas,'
and
all
and refugees
in
arms
border
these factors contribute to seriously
hamper economic development. Nevertheless,

May
in
2004, Sierra Leone held
its first
in
local elections
32 years, and the country continues
to
recover
395
World Atlas
Map
in
Sierra Leone
Data sources are provided at
Species
^-
Western ctiimpanzee observed 1980-1995
Western chimpanzee observed
after
1983
Estimated range
*iSS' Western chimpanzee
13"W
396
H-W
ttie
end of
ttiis
country profile
and economically from the
socially
devastated
latter
civil conflict.
infrastructure, however,
its
The
logging activity, however, and
and
the recent change
Sierra Leone remains at the bottom rank lout of
177 countries)
terms
in
Human
UN's 2004
of the
may
There have been multiple sightings
is
has
great ape found
of
widespread
in
Sierra Leone
chimpanzee [Pan troglodytes
the western
It
in
sparse
but
verus].
distribution,
occupying habitats ranging from young secondary
primary
to
woodlands, and logged

be found
south,
Outamba
Chimpanzees can
'
in
the east and
in
In
as an
now have
medical research
viable populations
hunting and agriculture
and so recent information on
Leone
scarce.
is
chimpanzees
of
2 000 individuals.'^
has decreased
is
It
at
tion
from tree
Loma Mountains
Hills Forest
Non-Hunting
Hills
which add up
Forest
a total area
to
of
the early 198Ds estimated
in
be
and-burn agriculture have resulted

and
the Kilimi section of
living in
of
soil exhaustion.
remains, and that there
of
chimpanzees
confirmed." A group
in
of
this area
has recently been
was
27 individuals
sighted
in
Tenkere, Outamba,' with a total population estimate

for
Outamba being 200-300.
park could
600-700
support a
It
is
thought that the
potential
population
of
The three Gola reserves IGola

and the largest, Gola North!
They
exploitation
in
were
lie
East, Gola West,

in
designated
the 1920s, but
the Eastern
timber
for
now comprise
the
deforestation
is
an average
360 km^ or 2.9 percent.^ Plantations
cash crops are replacing the remaining areas
and otherwise reducing the area
tat that
may
Eastern
in
of habi-
be suitable for chimpanzees.
Province,
In
the
diamond mining has resulted
large-scale modification of terrestrial habitat,
including forest clearance.

Sierra Leone
is
have been small areas
of
Although the overall trend
one
of forest loss, there
forest regrowth
due
to
in
reduced habitation during
the war; around the internal refugee
was heavy resource

forest
exploitation.
The
camps
total
there
area of
estimated by the Darwin-funded Habitat
Audit Project
individuals.'
Province.
of
in
estimated that 10 550 km^
is
It
of forest
the Outamba-Kilimi NP,' and the continued pre-
sence
and the pet trade.
pressure on the environment. Overharvesting of
Non-Hunting Forest Reserve, Kambui
to
cutting, hunting,
timber, the expansion of cattle grazing, and slash-
the proposed Lake Sonfon NP,
study
becoming
inadequate protec-
to
Rapid population growth has led to increasing
annual loss
835kml' A
The remaining
areas.
and declining due
isolated,'
of forest
its
chimpanzee presence mentioned
protected
populations outside these areas are
Island Wildlife Sanctuary, the Gola Forest Reserves,
49-60 individuals
of
in
Western Area Non-Hunting Forest Reserve, Tiwai
of
sig-
sign of
Surveys and
since then.
Outamba-Kilimi National Park INPI, the
and Tingi
little
found.'
records
above are
about
anecdotal data show that historically, chimpanzees
all
may have
the area
believed that the population
size
in
in
1990s,
early
the
In
chimpanzee numbers were estimated
Reserve,
in
THREATS
All the
has limited
conflict
of
numbers
the distribution and
Reserve,
there were scrapped after
facility
international protest. Later surveys confirmed that
thought,
is
It
the
important area for chimpanzees. Plans for a bio-
was
to researchers,
in
was estimated to be sparse.

Loma Mountains were proposed
1980s, the
existence
The recent history
occurred
the Gola reserves were minimal, and
in
the population
the woodland-savanna ecosystem of
or offer suitable habitats.'
Sierra
vations
reduced the population and
however, that few sites
access
young forest and 38
nificantly
the
of
the north of the country.
in
in
old forest in the mid-1980s. ^ At that time, obser-
Loma and
montane vegetation
and
Tingi,
forests.'
lowland rain forests
in
in
and savanna
gallery,
riverine,
forest,
profile!.
of solitary
the Tiwai Island Wildlife Sanctuary;
in
example, 22 individuals
for
Liberia the road
in
use (see the Liberia country
in
chimpanzees
Development Index."
The only species
not be
possible that with
is
it
government
in
is
about
000 km' lower than the
estimate of the Food and Agriculture Organization

of the United
of forest loss
is
almost
Nations (FAOl, but the estimated rate
between 1985-1986 and 2000-2003
identical, at
382 kmVyear." The Habitat
country's largest single area of lowland tropical rain
Audit Project identified no significant areas of cash
Gola North remains
crops and concluded that the few large plantations
"
forest.'-
Over 80 percent
of
unlogged, but satellite images reveal the existence

of a
logging road running from
reserve.
Liberia
The images showed no sign
into
of
the
major
in
Sierra
Leone had been abandoned
years, although
Satellite
images
some
of
rehabilitation
the
is
for
many
going on.
diamond mining areas
397
World Atlas
LEGISLATION
Sierra Leone
is a
signatory to the Convention on
International Trade
in
Endangered Species
of
Wild
Fauna and Flora (CITES), the Convention on

Biological Diversity,
and the International Tropical
Timber Agreement.
Ratification for all conventions
is
pending.' The national law protecting chimp-
anzees
Third
the
is
Schedule
the
of
Wildlife
Conservation Act of 1972. Chimpanzees are listed

as 'protected animals' and the young of the genus
Pan are
listed
as
specifically protected'.
same
the Fourth Schedule of the

individual
allowed two
is
These laws were under
act,
of
kills
Under
however, an
chimpanzees.'
revision at the time of
writing.
The long-running
recently caught
chimpanzee being
war and
civil
prevailing
engagement
insecurity that followed have inhibited
taken to the bushmeat
by the
market.
Sierra Leone. Earlier projects included surveys by
international conservation
community
in
Glyn Davies
and Baldwin
Teleki
dynamic landscape.
indicate a
are areas
In
the north, there
where the bush-fallow system
down, because the crop-forest rotation

be sustainable, but
to
system appears
insula there
coast,
is
increase
The
it.
chimpanzee-oriented conservation projects currently
the south and east the
uncontrolled development along the
is
is
being built that will further
above the new
forest on the hills
being cleared very rapidly.
Sierra Leone for at least 300 years. The
for Live
chimpanzees
boom
Led to a
in
for
demand
medical research overseas
the export of chimpanzees
in
the
Between 1973 and 1978, two wild animal
1V70s.
exporters from Sierra Leone are reported to have
shipped
overseas,"
582
chimpanzees
live
The
Netherlands.
happened
in
last
export
in
Rijswij(<,
to
Europe
1984.''
lacked any form
civil conflict,
protected areas
management, which
of
bushmeat marl<et
to sell
panzee meat
the
Tacugama Chimpanzee Sanctuary,

to the number of
which evolved as a response
chimpanzees kept as pets

it
in
the country; by June
held 62 rescued chimpanzees.' An acute
viral infection in
2004
at the sanctuary,
killed five infant
chimpanzees
prompting an emergency veteri-
nary intervention by
PASA
with the support of the
Zoological Society of London.'" The sanctuary
was
US government
with
later
awarded
which
a grant by the
around
to install electric fencing
its
15
km^
enclosure."
in
the southeast
is
Kenema
among those
primate meat. The quantity

in

The following recommendations follow the West
the
Action
and Conservation
Plan.''
of
chim-
bushmeat trade remains un-
Research
Basic data collection should
priority sites
be undertaken
in
such as the Gola Forest Reserves,
Outamba-Kilimi NP, Kuru
Hills
Forest Reserve,
are unaffordable to hunters, nets are used to trap
Loma Mountains, and Western Area

Non-Hunting Forest Reserve. A nationwide popu-
chimpanzees
lation
some
In
areas where firearms or their cartridges
for
bushmeat or
rural areas,
strength and
for the pet trade. In
chimpanzee bones are used
traditional medicine
398
is
Sierra Leone.
in
led to the
uncontrolled poaching of bushmeat.' The
known.
This
Pan
affiliated with the
During the recent
known
one sanctuary
is
African Sanctuary Alliance (PASA)
particular to Japan, the USA, and the
in
Biomedical Primate Research Centre

the
countries
to
underway'
There
2003
Chimpanzees have been hunted and trapped

in
Davies
breaking
be stable. On the Freetown Pen-
to
and a new road
developments
in
119841,
but there are no
(1989).
too short
is
is
Hardy
119811,
and Hanson-Alp
(19871,
vitality.
in
in
the belief that they increase
Tiwai Island,
census
of
chimpanzees should also be
initiated, partly to investigate sightings of
anzees near human settlements

of the country.
in
chimp-
various parts
Protection
as
The Outamba-Kilimi NP
is
believed
stronghold for chimpanzee conservation
Leone, so
it
should be a target
activities
through
promotion
of
Sierra
and
activities
Capacity building
Staff of the Wildlife
community programs. Foot and
strengthened
vehicular patrols should be undertaken periodically

in
protected
areas with viable
populations
chimpanzees. The Wildlife Conservation Act
of
should also be enhanced.
of
Population
Education
management
and maintaining corridors between
fragmented habitats that hold isolated populations
programs should be
Public awareness-raising
people
such
better trained. The
972
Establishing
areas.
Conservation Branch should be
numbers and
in
capacity of local nongovernmental organizations
should also be reviewed.
of
with at least one such program targeting
initiated,
wild
education.
tourism and conservation research,
possibly through
protects
and provides environmental
habitat,
conservation
of
antipoaching
in
houses confiscated apes,
it
chimpanzee
be the
to
the rural parts of chimpanzee range
in
Chimpanzee suppliers should be
will
be crucial
to
long-term survival
Leone's chimpanzees. Patches of natural
vegetation,
riparian
forests,
and mature bush-
fallows should all be considered as critical habitats
a focus of
International support should
initiatives.
provided to the
chimpanzees
of Sierra
when
be
Tacugama Chimpanzee Sanctuary
designing landscapes to
accommodate the
conservation needs of chimpanzees.
FURTHER READING
Alp, R. 119971 "Stepping-sticks"
and "seat-sticks": new types
American Journal o1 Primatology It^
Sierra Leone.
[11:
of tools
used by wild chimpanzees [Pan troglodytes]
in
45-52.
Fimbel, C. (1994) The relative use of abandoned farm clearings and old forest habitats by primates and a forest
antelope at Tiwai, Sierra Leone, West Africa. Biological Conservation 70
131:
277-286.
Sept, J.M., Brooks, G.E. (1994) Reports of chimpanzee natural history, including tool use,
century Sierra Leone. International Journal of Primatology\b

Squire, C.B. (2001) Sierra Leone's Biodiversity
and the
Civil
War
[61:
in
16th-century and 17th-
867-878.
Biodiversity Support
Program, Washington, DC.
http://www.worldwildlife.org/bsp/publication5/africa/176/titlepage.htm. Accessed June 12 2005.
MAP DATA SOURCES

Map
16.18
Chimpanzee data are based on the following sources with additional data by personal communication
from Davies,
G. [2005):
ButynskI, T.M. (20031 The chimpanzee

In:
Kormos,
R.,
Pan
Boesch, C, Bakarr,
troglodytes: taxonomy, distribution, abundance, and conservation status.

|v|.|.,
West African Cfiimpanzees: Status Survey and
Grubb,
P.,
Jones,
T.,
Davies, A., Edberg,
E.,
Starin, E., Hill, J. (1998)
Mammals
of Ghana. Sierra
Leone and The
Gambia. Trendrine Press, Zennor, UK.

Hanson-Alp,
R.,
Bakarr,
M.I.,
Lebbie,
A.,
Bangura,
K.l.
(2003) Sierra Leone.
In:
Kormos,
R.,
Boesch. C, Bakarr,
Butynski, T.M., eds. West African Chimpanzees: Status Survey and Conservation Action Plan.
M.I.,
lUCN/SSC Primate
Specialist Group. lUCN, Gland, Switzerland, pp. 77-87.
ACKNOWLEDGMENTS
This country study draws extensively on the Sierra Leone chapter from the
Status Survey and Conservation Action Plan.
Mohamed
Many thanks
(Ministry of Lands, Country Planning,
Wadsworth [Centre
for Ecology
and Hydrology, UK]
to Glyn
Forestry,
lUCN/SSC West African Chimpanzees:
Davies [Zoological Society of London], Hassan
and the Environment, Sierra Leone], and Richard
for their valuable
comments on
the draft of this section
COMPILER
Monitoring Centre
399
World Atlas
REPUBLIC OF SUDAN
Nigel Varty

Sudan, with an area
country
Africa.
in
tributaries,
the largest
is
dominated by the
is
It
per person. '' Agriculture
505 813 km',
of 2
and has borders with Egypt.
Nile
and
its
Republic
of
Ethiopia,
and
Congo IDRCI, Uganda, Kenya,
the
Eritrea,
It
on the Red Sea along

generally
its
in
complex process
in
the 1952 Revolution

1956.'
rule In
in
The
is
the east and west.
Sudan obtained independence from

Egyptian rule
of coastline
northeastern border, and
with mountains
flat
km
has over 800
Anglo-
joint
that
began with
Egypt and ended with self-
coup d'etat and military
first
Sudan has
616 270 km'
km'
6 410
of the country.'
permanent crops account
land and
percent of the
territory.' Tropical
1986-1989, both
terminated by military coups; the current form
government remains
Throughout
of
that of military dictatorship.'"
this history,
southern Sudan has
suffered from conflict between government forces
and
in
A peace agreement was signed
rebels.
1972, but
the
imposed
Islamic
throughout
law
Sharia
into
Army
Liberation
some
what
became
hundreds
internally
widespread famine.
which
People's
(SPLAl. The conflicts resulted
2 million deaths,
refugees and
Sudan
the
It
of
displaced
has been
thousands
and
independence,
to
northern,
maintain national
of the fighting
and
arm.
of the
Boma
the
at the
base
of
Plateau." Closed forest also occurs as
extensive galleries along rivers and

in
depressions
in
the southwest of Sudan.
and
people,
Muslim
clash
peoples
integrity. In fact,
is
divided.
international
the
Sudan

The eastern subspecies
much
populations
survive
with
found
in
finally
People's
Liberation
Movement ISPLM), to the negotiating table. Agreements have since been reached on a wide range of
is
of
forest areas on the border
as the Aloma Plateau near
only one recent (post-19831 record,
border region
the
Bengangai
Game
the
in
Reserve." This area
been a stronghold
civil
in
DRC and CAR, such

There
Sudan (west
the southwest of
in
Sudan. Relict
in
eastern chimpanzees are thought
of
Yei."''^
for
vicinity
of
of
Sudan has
southern forces during the
war. which has prevented safe access for
surveys.
By 2002. war-
pressure
chimpanzee. Pan
of the
troglodytes sctiweinfurthii, occurs
the V\/hite Nile! and
brought the government of Sudan and the SPLA's

political
in
forest in the Meridi district;
and Laboni forests
Lotti,
They
forests.
Aloma Plateau and
the Imatong Mountains; the Kinyeti Valley; and parts
to
has been factional and both northern
and southern opinion
weariness
the Talanga.
in
between southern, non-Muslim peoples seeking
seeking
Congo Basin
Yambio area; the Azza
of
complex struggle,
sometimes oversimplified as
is
the
were remobil-
country, southern resistance forces

izing
con-
terms gradually unraveled during
its
and by 1983. when the government
1970s,
is
CAR. DRC. and Uganda borders, representing the
include small areas on the
1965-1969 and
for only 7.1
moist forest
fined to a few small, scattered localities near the
then there have been only two periods of truly

in
although this includes
biomass resources are found
very northernmost
rule,
in
forests cover
the war-affected south of the country.'^ Arable
government followed within two years, and since
civilian
arid desert
Approximately 68 percent
of plantations.
of the country's forest

in
and
a tropical south
Woodland savannas and
the north.
and
GDP'
contributing 43 percent of
Chad,
Libya.
the Central African Republic (CAR), the Democratic
Sudan's most important
is
sector, employing 80 percent of the workforce
There has been no national census
panzees or recent studies
and
its
current
population
species
IS
status
estimated
to
of the
is
species
unknown.'
in
Sudan by
chimSudan,
^''
'^
The
be 200-AOO animals.' The
was already considered
endangered'
of
in
to
be
'highly
1988.'
issues that provide the political framework for a
THREATS
comprehensive peace agreement.
The population
of
Sudan, which consists
about 500 ethnic groups, was estimated

38.1
million
in
400
be
2003. with an annual growth rate of
2.7 percent.' In 2002. gross
was US$13.5
to
of
domestic product IGDPI
billion annually,
or less than
US$360
Sudan's
first
civil
war ended
in
1972, and peace
allowed considerable wildlife work to be done up

to
1983.
when
it
was brought
beginning of another,
much
to
longer
an end by the
civil
war Recent
surveys of wildlife resources of certain areas
of
<
AFRICA: Sudan
Map
in
Sudan
this
country profile
v*^'-
-M..
O
z
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,
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i,
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z
^
t
r
401
World Atlas
southern Sudan have begun only recently as part

rebuilding efforts
but there
no information on the rates
is
chimpanzee populations. Poaching

however,
rife,
of
anticipation of lasting peace,
in
known
is
been recorded, and there appears
be
to
marked contraction
in
have been a
to
range over the
of
in
the
abandonment and regeneration
bush and savanna forest
places.'"
50 years.
last
Disease
There was an outbreak
in
Hunting
Ebola hemorrhagic fever
of
Yambio, southern Sudan
in
May-June
2004.'*
Transmission from animals was not implicated
and there
this outbreak,
in
no evidence that the
is
disease has affected chimpanzees.
Hunting for bushmeat
considered a major threat
is
southern Sudan, and certainly threat-
to wildlife in
ens any chimpanzees remaining

Hunting
of the
consumption
Cessation of hostilities
war
the country.
The end
the
conflict
followed by large-scale deforestation as a result of
proliferation of firearms,
and the
expanded
greatly
because
and
sale
in
during
bushmeat
of
now
is
of the civil
the south
in
likely to
is
be
returning residents clearing land for agriculture,

cutting trees for timber to build houses,
and
for
commonplace.' Many former fighters and others
fuel to
now keep
security of the last four decades there has been
firearms, mainly AK-A7s, ostensibly to
The continuing lack
protect their livestock.

security has
made
livestock levels,
poaching
soldiers not to
Capture
meat. The Sudan People's
of
DRC)
young chimpanzees
officials
in
Chimpanzee Sanctuary
of
the
to
it
is
unknown
in
of the
Kenya, which had re-
in
Sudan
certainly being
used as
earlier
in
these
of
a conduit
markets.
food while waiting for crops to grow. This could put
any remaining chimpanzees
at risk.
LEGISLATION
Although national laws have been passed governing
most
wildlife conservation,
refer to the situation
applicable
in
the south, the regional government of
which has developed
its
own
set of legislation.
Chimpanzees are protected under the 1975

Wildlife Conservation
Regional
repealed
and Parks Act
Government,' but
(in
the
north)
The woodland and
forest
cover
of
Sudan was
estimated at 3A percent of the total land area at the

of
independence
forest or
.U
It
in
is
The
percent of the total
is
cleared
in
Sudan
rate of deforestation in the southern
areas thought to support chimpanzees
is
known, but these areas were strongholds
of the
not
Sudan Peoples Liberation Army during the
conflict
and both sides cleared forest around large
settle-
ments and
The
civil
Southern
of the
1986,
was
this
by the national Wildlife
military areas as a defensive
war drove many
rural areas, however,
and
measure.
millions of people from

killed
many more, which
989 (No.
for the creation of a
U of
number
reserve categories, and the 2002
989),
of forest-
Forestry and
Renewable Natural Resources Law.

The
1956, but had declined to
estimated that 9 590 km' of
each year,' although some authors give lower

figures."
in
Conservation and National Park Ordinance. Forest
which allows
7 percent by 2000.'
in
the north of the country and so have not been
under the national Forests Act
in-
reserves and other protected forests are covered
Habitat loss
time
meet
to
also likely to
crease significantly as farmers and refugees seek
Sweetwaters
Sudan was not the source
chimpanzees,
to illegal
intercepted
have been captured
ceived four other animals from

If
pet
In
in
southern Sudan.' Four
chimpanzees were taken
2003.
for the
Sudan.'
chimpanzees
10 young
Yambio
at
its
is
unable
endangered species.^
(thought likely to
origin
control
to
game
in-
and existing
south,
for
October 2003, Sudanese

of
the
demands. Hunting
trade has been identified as a threat
shipment
in
and wild animals are viewed as
kill
of
forestry activity
probably be
major species, and has directed
of
little
plantations would
difficult to
Movement has attempted
Liberation
of
meet domestic energy needs.' Given the
restock to previous
it
an alternative source
602
some
in
of decline of
areas where chimpanzees have
in
has resulted
tration
(WCGA)
is
General Adminis-
the national government agency
manage-
responsible for wildlife conservation and
ment and
the creation and
tected areas, although

of
the
country.
it
management
only operates
The management
in
of pro-
the north
forests,
of
including the creation of forest reserves,
comes
under the National Forestry Corporation, within the

Ministry of Agriculture and Forestry, but
vities
to
Its
acti-
have also been almost exclusively confined
the north
in
recent years due to the
The Higher Council
for
civil
war.
Environment and Natural
AFRICA: Sudan
Resources IHCENRI
agency
and was the lead
development
the
in
environment-
coordinates
related projects and programs,
National
the
of
Biodiversity Strategy and Action Plan.'^
As part
governmental organizations
Movement
Liberation
areas
sector-specific
of the
Sudan People
continue overseeing
will
southern Sudan, which
in
include both forestry and wildlife departments. This

joint
arrangement
expected
is
to
last for several
International
for
Development (USAIDI has been supporting work

determine the current status
through
Sudan
in
Sudan Transitional Assistance
its
program,' and aims
Rehabilitation (STAR!
to
of protected areas,
and biodiversity conservation
wildlife,
the peace process, the existing
of
US Agency
20 years. The
last
for
to
re-
create institutional capacity for conservation and
management
wise
natural resources.^
of
also been funding the
New Sudan
nongovernmental organization)
(a
USAID has
Wildlife Society
carry out
to
number
years during the transition period before the south
reconnaissance surveys
decides whether
parks, including Southern NP, considered the most
rejoin
it
separate from the north or to
to
within a single state.
International
Sudan
Convention
the
on
Biological
Diversity in 1995, the Convention on International
Trade
Endangered Species
in
Flora (CITES)
Desertification
UN
1982. the
in
in
to its size
still
Wild Fauna and
of
Convention
Combat
to
1995, the African Convention on
Conservation of Nature and Natural Resources

1973, and the World Heritage Convention
The country also participates
in
in
in
1974.
national
support chimpanzees,
to
coalition of organizations, including
Sudan Conservation
Society,
Sudan Conservation
Kenya
Sudan, but a
in
Born Free,
New
Wildlife Service,
Fund
Authority, International
Animal Welfare, and the Sweetwaters Chim-
for
panzee Sanctuary, has been helping with the rescue
and ongoing care
orphaned chimpanzees from
of
southern Sudan.
UNESCO's Man
and Biosphere (MABl Programme, under which two

Biosphere Reserves (the Binder
of
and remoteness.'
There are no ape sanctuaries
agreements
ratified
protected area
likely
due
in
NP and
the
Radom

There
is
no specific conservation action plan for
chimpanzees
NPl have been designated.
Sudan, although there are some
in
published recommendations dealing with the areas
Protected areas
in
There are three main protected-area categories
in
Sudan: national parks (nine terrestrial and two
game
marine];
reserves
(faunall
(221;
and sanc-
tuaries (including three wildlife sanctuaries!,
well as several other types of protected area.'
According
to the Ministry of Agriculture
protected areas
U percent
as
^
"
and Forests,
Sudan cover an estimated
in
of the country."
There are also many
which chimpanzees are thought
Although there
government
are
left
many
of
because records were

civil
these reserves are lacking

lost or
war Most parks and
destroyed during the

protected areas are
Chimpanzees are thought

and the Bengangai
to
occur
no recent information on the occurrence
panzees
torical
at
these
range
in
sites. Plotting of their
to the
in
of
chim-
recent his-
the Lantoto NP.
long-running
civil
war
Sudan there have been no conservation

the eastern chimpanzee.
detailed
the
Sudan suggests eastern chim-
panzees may also occur
Due
in
Game
Reserves,'
Reserve.'" "' There is
Game
to
occur
to
Sudanese and
the southern
in practice,
human
their
protected areas and re-
most protected areas
settlement, cultivation, and
livestock grazing. There are also
few land-use plans,
so surveying wildlife areas and preparing wildlife-
management
plans merit priority attention.'
'"
There have been calls for the Sudan Peoples

Liberation
Movement
to
consider making "unequiv-
ocal and very public pronouncements of absolute

prohibitions against the hunting of elephant, rhino,
and chimpanzee".' Incentives have been proposed
inadequately staffed and financed.
Mbarizunga and Bire Kpatuos
open
still
reported to be considerable
to re-establish
constitute them,^
forest reserves in Sudan, particularly in the south,
but data on
among
support
is
in
southern
projects for
Nor have there been any
chimpanzee research projects within the
to
encourage armed groups
firearms. There
is
to
decommission
tegy for the conservation and sustainable
ment
of
the
country's forests and
particularly for high forest areas
and capacity building and

ening
in
their
also the need to develop a stra-
in
manage-
woodlands,
the far south,
institutional strength-
natural resource planning and strategy'
Given that Sudan has just been through
than 30 years of
civil
more
war, with millions killed or
displaced, and with very few financial resources,

wildlife conservation
is
unlikely to be a high priority
403
World Atlas
management, taking
immense human needs
tor either the northern or the southern governnnent,
conservation and
and outside agencies and nongovernmental orga-
account
most
nizations are expected to provide

for the foreseeable future.
is
funding
country, particularly
The over-arching
priority
tional conservation Interest In
emerges, determining the status
and developing a
the
of Its
to continue to survey present conditions as
tions
of
of wildlife
prioritized
unreasonable
peace
In
the
In
the south. Given the Interna-
chimpanzees,
It
not
Is
expect donor support for any areas
to
that are found to harbor
popula-
plan for their
careful
remnant populations
of
these great apes.
FURTHER READING
Catterson,
and Opportunities Assessment. USAID Integrated Strategic Plan
(20031 Environmental Threats
T.
Sudan 2003-2005. USAID/REDSO/NPC and the USAID Sudan Task
the
in
Force, Washington, DC. http;//vmvw.usaid.gov/
locations/sub-saharanafrica/sudan/sudanjspâ^.pdf. Accessed June 12 2005.

El
Moghraby,
A.I.
12001) State of the environment
in
Sudan.
In:
UNEP EIA
Training Resource Manual.
UNEP.
pp. 27-36.
vww.unep.ch/etu/publicatlons/111%2027%20to%2036.pdf. Accessed June 12 2005.
HCENR
and Action
(2000) The Sudan's National Biodiversity Strategy
Plan. Higher Council for Environment
and
Natural Resources, Ministry of Environment and Tourism, and lUCN with support from UNDP, Khartoum.
http://wvm/.biodivorg/doc/world/sd/sd-nb5ap-01-pl-en,pdtto http://vi(ww.biodiv.org/doc/world/sd/sd-nbsap-01p7-en.pdf. Accessed
June 12 2005.
MAP DATA SOURCES

Map
16.19
Stevens,
E.F.,
In:
following source:
Beck, B.B., Stoinski, TS., Hutchlns, M., Maple,
Arluke, A., eds. Great Apes
and Humans: The
TL,
Norton,
B.,
Rowan,
A,,

ACKNOWLEDGMENTS
Many thanks
Thomas Catterson (USAID Sudan
to
Natural Resources
&
ations are recorded." The main forest areas are
in
International
Environmental Management! for his valuable comments on the draft
Consultant,
Forestry,
of this section.
AUTHOR
Nigel Varty.
UNEP World
REPUBLIC OF UGANDA
Nigel Varty

The Republic
of
Uganda
lies
on the Equator
in
the west of the country.
East-Central Africa, mostly between the northern
and western shores

Valley.
Its
Victoria
Albert.
territory
Lake Victoria and the
includes nearly half of Lake
who was overthrown In a coup d'etat in 1971."" The

new dictatorship, led by Idl Amin, persecuted and
of
Lakes Edward and
Hence land occupies only about 85 percent

its total
swamps amount
to
area of 236 OiO
another
kml
Large
percent of total area,
while forests cover 41 900 km^ of Uganda, representing about 21

forest
iOi
is
In
1962, with an elected prime minister IMIlton Obotel,
and substantial parts
(199 710 km'l of
Uganda became independent from the UK
Rift
of
percent of the land area. Most
natural woodland, but 430
km'
of plant-
expelled
members of the Ugandan Asian community
and the country's

of
intellectuals.
Tanzania invaded Uganda
of exiled
The United Republic

979 with the support
Ugandan groups. AmIn was overthrown,
and elections
The
In
validity
of
In
1980 returned Obote
the elections
to
power
was challenged
by
AFRICA: Uganda
Army
Yoweri Museveni's National Resistance
which
and eventually won a
initiated
Museveni as president
installing
charge
country
of a
in
which
1.5 million
war,
He took
1986.
in
INRAI.
civil
people had
maimed in war, 2 million were refugees, and the economy was in ruins. Considerable
progress has since been made in restoring peace
been
and
killed or
rebuilding infrastructure and
in
Museveni was elected
civil Institutions.
to office in the elections of
but these are part of a larger VIrunga population.
was estimated to number 380 individuals In

among the contiguous Volcanoes
This
2003, distributed
NP
ing
now
is
the spiritualist
is
ILRAI, which, despite
been recorded
stated aim of wish-
its
atrocities since 1986.
It
the Ten
to
believed to have abducted
is
20 000 children over the years, using them as
expendable fighters, and the number

people
living
camps
refugee
In
estimated at about
in
of displaced
the north
supported by Sudan, which suspected Uganda

supporting
own
its
Sudan country
the
rebel group
but
profile],
ceasefires and
attempts
failed
of
1999, both coun-
in
support the
to
A succession
insurgents.
respective
primatologlsts consider
of
western Uganda,' having
different forest blocks since
21
In
upwards, although populations have been
declining. In 1979,
It
was thought
of
breeding females."
was estimated
1989.
In
This
was
down
revised
to 3
at
broken
of
negotiations
1997.'^
In
In
was US$6
product IGDPl
growth rate
billion,
per person,'^ and the country's
or about
Human
of 177."
the territory and agriculture
tant sector of the
of
However, the
The mountain
stable
population),^'
populations appear
gorilla
or Increasing
(Bwindl)^'
Is
but they and their habitat are

a variety of
drastically
US$270
Develop-
It
is
estimated that 910 km'
of forest Is cleared in
Uganda annually, representing about
2 percent of
the total each year during the period 1990-2000."
percent
Tropical high forest covered about 13.7 percent of
the most impor-
the land area
and accounting
for
in
1900, but had been reduced to about
percent by 2000.' Illegal logging, generally

pit
sawing,
is
widespread
Uganda, particularly
of
species
such as the mahoganies
Entandroptiragma?'' Charcoal burning
[Gorilla beringei beringei]
and the eastern chimpanzee [Pan troglodytes
NP
in
in
the VIrunga
southwest
DRC, and
away.
to
Uganda. Mountain gorillas
only two localities:
of the
in
Only
In
the Mgahlnga Gorilla
mountains
at
country bordering
the
Bwindl Impenetrable NP, about 25

12
be present
In
km
mountain gorillas were thought

the Mgahinga Gorilla
NP
by 1998,'
some
timber
Khaya and
is
forest reserves,
also hav-
such as
Kasyoha-Kltomi and Kallnzu. Indeed, 90 percent

of
the
fuel
nation's
energy source
of
derived
from
critical
chimpanzee
reserves by sur-
forest
rounding farming communities

of
is
wood."
Encroachment
extreme
Rwanda and
an impact on
the forest
in
for valuable
reserves
ing
occur
recent
in
Habitat loss
schweinfurttiii] occur in
still
causes. Chimpanzee
The mountain
gorilla
be
nearly
involving
force
to
(VIrunga
slightly
decades,"^' probably as a result of habitat loss.
work
of the
of GDP.'
7001.''^
THREATS
3.6
80 percent
based on
latest estimate,
about 4 950 (range 4 000-5
economy, employing more than
43 percent
chimpanzee
Permanent
crops and arable land account for 34.1

of
of
2003, the gross domestic
ment Index rank was U6th out
in
data gathered between 1994 and 2002, gives a figure
numbers have declined
The 2002 census estimated Uganda's popumillion, with a
300
that
Uganda."
In
population and habitat viability assessment (PHVA)
conflict continues.
lUl
it
there were about A 000 chimpanzees
under threat from
lation at
were
that there
which 750 were
fewer than 3 000 Individuals,
between the government and the LRA mean that the
3 percent per year.'"""
have slowly been re-
1994.^^ Population estimates
vised
the south (see also
In
reached an agreement not
tries
is
The LRA had been
1.6 million.'^
woodlands
and east
Commandments, has been committing widespread
some
some
although
gorillas,
forests and
Resistance
Lord's
govern the country according
to
Hence, this park
the Bwindl gorillas to be a separate subspecies."
relatively
troubled by violent Insurgency'^ The main
still
Army
Uganda
half of
but large areas of the north
rebel group
the southern section of the
DRC.'' There are about 320 gorillas
Impenetrable NP.
Bwindl
In
in
The eastern chimpanzee occurs across the
The southern
are
NP
contains just under half the world's mountain
1996 and 2001.
peaceful,
Rwanda and
In
VIrunga
Is
habitat,
reducing the area

but the
conversion of forest to agriculture
is
greatest
currently
taking place outside forest reserves, with large-
/;05
World Atlas
Map
in
Uganda
country profile
34*E/
3ZE
.^306,
this
Aro?,
'-,
DEMOCRATIC
"0^
REPU BL C OF
-^
"^
.i,
.
TTnP
I
V*.-,
-G"'"'
*^
THE CONGO
Semliki
kanyawara
Rwenzori
-,'
'fwtountains/
-T,'' "d
NP & WHSX-'
i
O'KT.
\rKamp,
f<y
lYANCHu
r7
:)
KENYA
}y
Bigodi Wetland SancSiary
<..!.-
J^
.,
i Kyambura WR
\ r^gamba
Island
jij.Chimpanzee Sanctuary
\-
Kasyoha-Kitomi FR
>'
EilwurJ
Queen
l-i/^-!f-\>''
y^Kalinzu
Mbarara
Elizabeth Npi-
FR ^__,,x^
/ Maramagambo
Masaka.
Wr
.Ss^*^-'
^
'^
* l,'S'!">!luc
^^P&WHS
^j^jj-,-\^
ke V i c t o
Species
/Mqezi
^-Sl'*'^
.^^^
UNITEDREPUBLIC
OF TANZANIA
Eastern chimpanzee obseived 1996-2003

Eastern lowland
Mountain
gorilla
gorilla
Igahinga
Gorilla
NP
Confirmed range
RWANDA
Mountain
gorilla
Estimated range
^'.Vjl
Eastern chimpanzee
Eastern lowland
3(r^
406
gorilla
2=3
AFRICA: Uganda
scale plantings of tobacco and
sugar cane and tobacco
district,
and tea around forests
in
Hoima
Masindi
district,
cocoa
in
Kabarole
in
flow.
chimpanzees, hence preventing gene
Comparison
of satellite
mid-1980s and 2000-2001

800 km^
chimpanzee
of
showed
appear
that
about
was
lost in
Other threats include mining, particularly
which
most recently
forest,
used
for
manufacture
the
in
NP
have been reported
from scabies [Sarcoptes
suffering
known
is
five juveniles
badly affected."'
infections
thought
electronic goods.^^
domestic animals
living
rates
with
is
in
of
Hunters
Uganda
in
sometimes
still
gorillas,^ but regular monitoring
reduced the incidence
most
part,
surveyed
recent years. Ugandans,
in
do not eat chimpanzees. Although
Plumptre
by
mountain
kill
and patrolling has
signs of hunting were recorded
is
the forests
all
in
and
NP
Ruwenzori Mountains
the
colleagues,
where
the only one
chimpanzees are regularly hunted

is
DRC, who do not share the same cultural taboos,

could have a serious impact on remaining popu-
and chimpanzees are
lations. In addition, gorillas
affected by snares set for animals such as duikers
and bush
pigs.
Snared hands or
ously mutilated or even
and the resulting
lost,
More than
tions can be fatal.''
and up
of
forests
have snare-related
in
Budongo's 'Sonso' habituated group,

deaths are suspected
to
be
"
injuries.'''
if
"
'"
In
two
at least
killed by
crops around forest reserves and
national parks have been raided, particularly
if
cash
crops are involved. There are reports of spears and
bows-and-arrows being used
Bugoma
for taking
sugar cane
chimpanzees
been found
The
for
body parts
in
animals
kill
Budongo
at
Evidence of hunting
for ritual
at
for
of
use has also
many
young apes
is
less
to
be sold as pets
common
other countries,
but
in
Uganda
many young
chimpanzees are smuggled over the border from

DRC. There has been a recent increase
ber
of
chimpanzees confiscated
in
forest.'"
or
High
the Bwindi gorillas." The

is
particularly worrying
gorilla
demand and
for
and chimpanzee
tourism
in
Uganda,
the potential financial
The enforcement
regulations put
in
place to reduce the risk of disease transmission
is
benefits.'
'
of
weak.
still fairly
LEGISLATION
National and international law
Uganda has
ratified or
the
acceded
num-
(three to four a
the Convention
to
on Biological Diversity 119931, the Convention on

International Trade
in
Endangered Species
Fauna and Flora ICITESI

to
Combat
UN
(1991), the
of
Wild
Convention
Desertification 11997), the Convention on
Migratory Species (2000), the African Convention on
Conservation
(19771,
of
Nature and Natural Resources
and the World Heritage Convention
Two World Heritage
Sites have
(1987).
been inscribed - the
NPand Ruwenzori Mountains
Bwindi Impenetrable
in
UNESCO's Man
and Biosphere (MABI Programme, and Queen
NP
Elizabeth
has been designated as a Biosphere
Reserve.
Chimpanzees are protected by

national and
restricted.^'
international trade
is
law,
and
regulated or
There are two main statutes that most
concern protection of great apes

Forests Act 119641 and the
(1996), respectively
at Kalinzu Forest Reserve.''^
collection of
or for entertainment
than
and
cacao,
fields.''^
to
of the
humans
Cryptosporidium parvum
in
groups are being habituated

to tourist
this
NP. The country also participates
a result of snaring.
Chimpanzees have been hunted and

local people
25
to
Budongo
Kibale and
percent
infec-
half the population
(nine adults) in the Kalinzu forest,
chimpanzees
can be seri-
feet
near
human-ape disease risk

as increasing numbers of
due
One
There
for meat.'*
increasing concern, however, that refugees from
raiding
infection
of
The source
'
have been
to
have also been reported
Hunting
for the
is
scabiei].
have died from the disease, with
to
semiconductors for mobile phones and other
coltan,
which
of
or their livestock. Habituated gorillas
Bwindi Impenetrable
infant
in gorilla
Uganda, some
in
have been contracted from close contact
to
humans
with
in
Kasyoha-Kitomi
diseases have been reported
of
and chimpanzee groups
this period.^'
the
of
Disease
A number
imagery between the
habitat forest
withdrawal
the
of
Uganda's military presence from DRC.'*
This
district.
reduces forest connectivity and dispersal opportunities for
because
possibly
year),
Forestry Authority
in
Uganda
Uganda
- the
Wildlife Statute
executed through the National

(NFA)
(formerly the
Forest
Department) and the Uganda Wildlife Authority

(UWAI,
in
Industry.
the
Ministry of Tourism, Trade,
The Uganda
and
Wildlife Statute provides for
the establishment of wildlife conservation areas,
which
fall
under two categories:
wildlife protected
407
World Atlas
areas (national parks or wildlife reserves) and
management areas
wildlife
and community
2004,
In
sanctuaries
(wildlife
criticized for negotiating
Changsha
the transfer of three chimpanzees to

Zoological Garden
Injunction
In
was Issued
China as a
at the
now
is
put
state."
gift of
An
venting the transfer until a hearing
In
May
was murdered
In
of
remaining fragment
The
Uganda.
where lowland moist
country.^^
There are three main categories
protected area
In
Uganda: national parks
wildlife reserves (10 sites],

sites).
of
(10 sites),
and forest reserves (710
Forest reserves represent close to 50 percent
While most are less
of the protected land cover''
than 10 km^
in
some
area,
sented by just seven
Budongo,
Bugoma,
2 600
'central'
Kallnzu,
km^ are repre-
forest
reserves;
Kasyoha-KltomI,
Maramagambo, and Sango Bay

Reserves.'' '^ Some forest reserves include
Mabira,
tions of exotic species
Mgahlnga
Forest
planta-
such as eucalyptus.''
NP
Gorilla
is
Bwindl than
In
rounded by one
In
In
In
an undis-
less
human
main
the
gorillas have
Bwindl Impenetrable Forest Conservation Trust and
program
to
share tourism revenues with local
communities have been established." There have,
been conflicts over revenue-sharing
KIbale
NP
(780 km')
Is
the most important
protected area for chimpanzees
Uganda.
in
was completely
set-
25 percent of Uganda's chimpanzees.''
In
a favored gorilla habitat,
1991.
The
settlers
to
the park being gazetted
were evicted and the eco-
by
tourists. To sustain the park in the long term, the
supports a population
and cultivated prior
In
sur-
population visited
gorilla
was
once
is
populations
population growth. The open woodland, which
tled
Among
bamboo
and surrounding lands are
Rwanda, the Bwindl
in
become
the park
human
is
East Africa
and has been very successful. Since
April 1993
the genocide
however,
to
much
and
In
Intensely cultivated. Gorilla-based tourism started
arrangements."
of
Is
densest
of the
Africa (100-450/km'),
national park, at only 34 km',' and harbors part of
has been habitat loss or modification due
valleys,
forests blend
in
extremely
is
the VIrunga region." Bwindl
the VIrunga population of mountain gorillas. The
major problem facing the conservation
terrain
the few large expanses of forest
other differences, there
the
is
natural forest
turbed continuum Into montane forests.
Uganda's smallest
(321 km')
of
and narrow
hills
Altogether Uganda's protected areas cover about
of
NP
Impenetrable
Bwindl
largest
rugged," with steep
or 14 percent of the land area of the
in
honey, water, and firewood.
one
km^
to
Pressures include hunting, grazing
Protected areas
33 000
began
that
the area
southwest
2004.
measures
place the
park,
beekeeping, and collection of bamboo,

''^
livestock,
request of the Green-
watch nongovernmental organization (NGOl, pre-
''
1994."
regenerating. Sucker, the warden
In
restore the
wildlife areas).
Uganda was
system
who had
around
of
It
430-1 530, about
'"'
" The
density of chimpanzees recorded here (2.32/km')

is
the highest of any forest surveyed
in
Africa.'^
Three chimpanzee communities are habituated:

Gordon Mider/IRF
Kanyawara and Ngogo
smalt township
market
in
Uganda, close
to ttie border with the
Box
Democratic Republic
park,
the Congo.
of
behavioral research
for
purposes, and Kanyanchu for tourism activities (see

4.6).
Forest covers nearly 60 percent of the
with
degraded
which
forest,
is
largely
secondary forest recovering from agricultural use
in
the southern sector, covering another 9 percent.'
Its
designation as a national park

Illegal logging
in
1993 has reduced
and agricultural encroachment, but
crop raiding by animals, including chimpanzees,
and the laying

still
of
snares
in
the forest by villagers,
contribute to strained relations between park
managers and
local communities.'^
tourism area
at
killed several
human
to conflict
These
Near the eco-
Kanyanchu, chimpanzees have

infants,
which has also
between the park and the
killings
led
local people.
were not carried out by habituated
chimpanzees.^'
Chimpanzees occur
(08
in
many
other protected
AFRICA: Uganda
areas
Uganda, including
in
NP, Rwenzori
Sennliki
Mountains NP, SemlikI and Kyambura Wildlife

Reserves, and at least
of
forest reserves.'^ Several
1 1
these areas combined are estimated to hold over
80 percent
0581 of the total population. These are
[A
Budongo Forest Reserve

Reserve
Forest
NP
Kibale
(500),
Reserve
Mountains
Kasyoha-Kitomi Forest
-129),
(1
Bugoma
NP
1639 animals),
Ruw/enzori
1628),
Kalinzu Forest Reserve (234), and
(4061,
Maramagambo
Queen Elizabeth NP
forest inside
(222).'^
Chimpanzees have also been recorded
Mount
Otzi Forest
the
in
border with Sudan." The remaining chimpanzees

are found
small numbers
in
between
pockets
particularly
these
the
in
Many
districts."
Bwindi Impenetrable
Reserve on Uganda's northern
of
communities are not
forest
isolated
in
larger
blocks,
forest
National Park.
Gordon Miller/IRF
Kibale
Uganda has
field
Budongo
these isolated chimpanzee
also had the additional effect of reducing forest
in
protected areas.
degradation.^
research on chimpanzees
long program of research
particularly
"''
Kibale forest.""
in
lUCN-The World Conservation Union
involved
also
several
in
and
conservation
development projects around major conservation
conservation
Budongo, ' with
in
forest, in collaboration with
Project (KCP) and
Forest Project (BFPl respectively These efforts have
a history of
1962
dating to
Chimpanzee
Hoima, Kibale, and Masindi
is
Research and
NP and Budongo
the Kibale
areas important for chimpanzees, including Kibale
NP and
Semliki NP. The Jane Goodall Institutes and
(WCSl have worked
Uganda
Research on mountain gorillas has been ongoing
with the
since the 1950s.' Following increased security
Department (now the National Forest Authorityl
Uganda
started
in
in
Reserve,
in
the 1990s, several other studies were
Budongo Forest Reserve, Semliki

Impenetrable NP, and
Bwindi
between
Mgahinga, and
gorillas
the
gorillas
Wildlife
panzee conservation, following
Kalinzu
the national
Bwindi
of
of the dietary differences
and chimpanzees
and
between
Bwindi Impenetrable
at
NP." As elsewhere, the commitment
of particular
individuals has led to the
development
long-term study
Uganda these include
Struhsaker,
sites.
In
Kanyawara; Mitani and Watts

in
Sonso and Budongo; Hunt
these
in
in
Ngogo; Reynolds
in
Semliki; Stanford
Bwindi; and Hashimoto and Furuichi
National conservation
of
and Wrangham
Isabirye-Basuta,
research
in
facilities
in
Kalinzu.
include
to
Technology, and
University's field
Kibale NP, which
ever
field
In
Mbarara University
is
based
station at
is
ation
some
of
of the longest
addition to conservation
have been active
Uganda.
Institutes (JGI)
work promoted
number
in
their joint
work on
to resolve
issues through
human-chimpanzee
collaboration
on snare-
removal programs, alternative income-generating
schemes, and sustainable development issues.
It
also facilitates environmental education programs

for
primary-school children, particularly those

near chimpanzee habitats, and plans
living
develop the
same schemes
for
and adults. The Jane Goodall

developing
groups
Roots
for local
&
to
secondary schools
Institute
is
also
Shoots' environmental action
communities.
Other conservation and development projects

include activities of
WWF-The
Global Conservation
Organization around Ruwenzori Mountains NP, and
CARE
International's
Impenetrable
WCS
around
activities
NP and Mgahinga
also support the
Forest Conservation, and
Institute
WWF,
Bwindi
Gorilla NP.
of
WWF
Tropical
together with the
of interna-
African Wildlife Foundation (AWFl and Fauna and
great ape conserv-
Flora International (FFIl, supports the International
partnership of the Jane Goodall
and the Uganda
to
chimpanzee survey."
communities
conflict
and
studies of primates.
NGOs
in
Science and
and Makerere
Kanyawara, north
the base for
by international researchers, a
tional
of
at Ruhija,
Forest
The Jane Goodall Institute-Uganda works with

local
the Institute of Tropical Forest Conservation (ITFCI,
which belongs
Authority and
develop a national species action plan for chim-
Forest Reserve.'^ Comparative studies have been
ongoing
Wildlife
Wildlife Authority
has been carrying out snare-removal programs
in
Gorilla
Conservation
Programme
(IGCPl,
which
runs and supports projects at Mgahinga Gorilla
NP
and Bwindi Impenetrable NP.
409
World Atlas
Sanctuaries
The Uganda
holding
Wildlife Education Centre
for
facility
Uganda, and
home
also
is
Ngamba
established
1998
in
community
to a
in
of eight
other animals.
a variety of
to care for confiscated
orphan
managed
by the
Uganda.
in
animals
Chimpanzee Sanctuary was
Island
chimpanzees
the official
is
confiscated
all
Chimpanzee Sanctuary &

Trust (CSWCT), which
is
It
Wildlife
Conservation
a collaboration of six inter-
is
national trustees: the Born Free Foundation, the
staff at Kibali National
Fund
International
Park.
Animal Welfare, the Jane
for
Julia Lloyd
Goodall Institutes, the Environmental Conservation
Tourism
Trust of Uganda, the
Nature-oriented tourism, including gorilla and
Centre, and the
chimpanzee watching, has been developed

eral sites
NP
NP
Uganda, including
in
Mgahinga
(gorillas),
Bviîndi
NP
Gorilla
at sev-
Impenetrable
(gorillas!, Kibale
(chimpanzees), Budongo Forest Reserve (chim-
influx of
but,
following an
orphans, the sanctuary
now
cares for over
and
tourism
gorilla
of
Uganda commissioned by IGCP estimated
in
that in
located
is
Entebbe and Isinga
in
Queen Elizabeth NP,
35 chimpanzees.
An economic evaluation
Ngamba
Lake
in
Island
Victoria,
is
0.A5
23
for the
chimpanzees, but as the forest

to sustain this population, the
animals are provisioned. The sanctuary
million
in
in
government
sales, contributed US$'i.77
and US$6.93 million
taxes,
to
the national economy.'" Gorilla watching at Bwindi
Impenetrable
income
of
NP
generated up
is
is
for
day and overnight
300
visitors per
&
Wildlife
visits,
attracting
Conservation Trust works with and
education, health, sanitation, and 'sustainable living
on Lake
visits
tourism
are recovering and the
opened
Bwindi
at
eight tourists and a
in
new
July 2002.
gorilla
Uganda
considered one of the best countries to see
chimpanzees
the wild," and Kibale
in
is
Victoria'
programs, micro-finance and loan
schemes, and provides casual labor and permanent

employment. The
trust also runs training
the capacity of
to build
a parti-

In
ape tourism
ape conservation;" great ape
1999,
Jane Goodall
the
tourism can have potentially damaging impacts
year program,
on the animals themselves as well as on their
Wildlife
One
development
issue, for
extensive
of
damage
improves access
example,
to
the
trail
forest
for hunters.
is
whether the
systems does
understory and
Research
to clarify the
lation
in
be noted that
trail
systems that are frequented
by tourists, researchers, and guards are likely to
deter poachers, potentially offsetting any negative
impacts
of trail cutting.
a four
Uganda
Uganda
Forest
and
the
viability
is
built
on
popu-
assessment undertaken
1997 by the Conservation Breeding Specialist
develop a
can
and the
evaluate the current status of
to
and habitat
and chimpanzees
it
Institutes
commenced
chimpanzees.'^ This program
Group
underway.'"" Meanwhile,
in
Authority
Department,
significance of such impacts for mountain gorillas

is
captive
in
chimpanzee management.
raised, however, about the net benefits of great

for great
programs
Ugandan nationals
cularly successful site for this. Questions have been
excessive
more than
month. The Chimpanzee Sanctuary
DRC murdered
Wildlife Authority
habitats. ^^
run as an
is
a popular visitor destination
March 1999, when Rwandan rebels
in
park warden. However,
for
education center and
benefits neighboring island communities through
an incident
Uganda
50 percent of the
not large
Uganda's national park system before
traveling from
group
to
area
enough
environment
exchange earnings
about US$7.7 million, gener-
in
southeast
Entebbe. The island provides a semi-natural
of
IS
of
km'
km
199^-1999, gorilla tourism attracted net foreign
ated US$15.4 million
410
Education Centre
Island of
panzees), Semliki Wildlife Reserve [chimpanzees),
Reserve (chimpanzees).
Education
Wildlife
Wildlife Society Originally,
chimpanzees were relocated from the Uganda
19
Wildlife
and Kyambura
Wildlife
Uganda
Uganda
of
lUCN." The results have been used

five
year national action plan," which
to
is
currently being reviewed by the Minister of Tourism,
Trade, and Industry.
The
five
action plan"
year goal of the national chimpanzee

is:
"To strengthen the protection of
AFRICA: Uganda
chimpanzees and enhance the

lations
corridors.
tives
popu-
viability of
major forest blocks by establisfiing
in
In
"
order to achieve this goal,
objec-
six
were defined:
ment and implementation

policies to
of
environmental
promote chimpanzee conservation;
develop a health monitoring
program
fund improvements
reduce fragmentation and loss
of
key chim-
available
public health facilities
in
communities
to
in
near
areas
panzee habitats;
tourism and research
reduce conflict betvêen local communities
evaluate and
and chimpanzees;
panzee research and tourism guidelines;
promote awareness
reduce levels
injuries,
of
chimpanzee values;
affects
responsibility
where
chimpanzees;
risk
to establish
health
guidelines for tourists visiting chimpanzees;
Uganda and enhance

[e.g.
disease transmission
of
develop standardized chim-
improve chimpanzee tourism experience
and the pet trade;
minimize the
sites;
fund and develop a project
human-caused deaths and
of
enhance corporate social

it
tor
chimpanzees;
between people and chimpanzees.
improve
its
facilities,
in
product marketing
invest in habituation,
improve guiding experience);

evaluate the impact of ecotourism on chim-
panzees, local people, and the environment.

Several projects and activities were identified for
each
The lUCN/SSC Primate Specialist Groups African
objective, including:
Primates: Status Survey and Conservation Action

develop a monitoring system that
allow
Department (now the National
Forest
the
will
Forest Authority] to detect illegal activities

quickly and to bring
them under
in
Bugoma, Budongo, and Kasyoha-Kitomi;

awareness
the
and protection
things] continued monitoring
Virunga and Bwindi gorillas, studies
to
of the
assess the
most appropriate tourism strategy and arrange-
control;
develop and fund snare-removal projects
raise
recommends (among other
additionally
Plan""
ments
for the
two areas, and improved programs
for conservation
awareness.
law enforcement
of
agencies (customs, police,
etc.),
international
Research and education
human
organizations lUN, diplomatic corps, airline
Research
companies,
Interpol], for
disturbance on gorillas and other wildlife at Bwindi
ing posters
and
example by develop-
leaflets for distribution,
ensure that relevant knowledge

ated
in
is
and
incorpor-
law enforcement training;
work with
private
landowners
investigate
have
if
and
what attributes corridors must

to
be able
to
move
design chimpanzee-proof beehives for local
communities around
forest blocks;
people
and development projects with
living
of
the
effects
of
concern include tourism
edges
of the
park for beekeeping,
collection of non-timber forest products, etc.^'
The development
of
for schools in the Biiso,
Pakanyi
(south)
in
conservation-education
identified
as a particular priority
Budongo, Kijunjuba, and
areas around
Budongo Forest
the Mabale area around
Bugoma
Forest Reserve.'
Julia Lloyd
develop and fund community conservation,

education,
Impenetrable NP. Areas
Reserve, and
them;
into
and the multiple-use zones that have been estab-
programs has been
forests);
chimpanzees are
freely along
needed
lished around the
to identify
maintain habitat corridors (particularly around
Bugoma and Budongo
is
Tourism
at Kibali
National Park.
around Budongo, Bugoma, and
Kasyoha-Kitomi forests;
implement
a national environmental
aware-
ness campaign, with a focus on great apes,

especially for primary schoolchildren, but also
for
secondary schoolchildren, and incorporate
within the national curriculum;
improve environmental awareness among

corporations, which will lead to the develop-
411
World Atlas
FURTHER READING
Edroma,
Rosen,
E..
Assessment
N., Miller, P.,
eds 119971 Conserving the Chimpanzees of Uganda: Population and Habitat
lUCN/SSC Conservation Breeding
for Par) troglodytes schweinfurthir
Viability
Specialist Group, Apple Valley,
Minnesota.
Howard, P.C, Davenport,
Mupada,
T.R.B., Kigenyi, F.W/., Visl<anic, R. Baltzer, M.C., Dicl(inson, C.J.,
Protected area planning
E. 120001
Conservation Biology
McNeilage,
|3I:
OryxaS
A.J.,
Reynolds,
2.
R.A.,
ID: 39-47.
Cox,
Mugume,
D.,
Institute.
S. 120031
Wildlife Conservation Society,
120051 The
V.
Struhsaker,
Matthews,
858-875.
Conservation Society/Jane Goodall
Series
J.,
Arnold, M., Nkuuta, D. 12003) Conservation Action Plan for Uganda's Chimpanzees 2003-2008. Wildlife
A.J.,
Ptumptre,
Lwanga,
the tropics: Uganda's national system of forest nature reserves.
Plunnptre, A. J., Brock-Doyle, A.. Weber, A. (20011 Bwindi Impenetrable National Park: gorilla census
A.,
1997.
Plumptre,
in
T.T. II
The Status of Chimpanzees
New
in
Uganda. Albertine
Rift
Technical Report
York.
Chimpanzees of the Budongo
Forest. Oxford University Press, Oxford.
9971 Ecology of an African Rain Forest: Logging in Kibale
and the
Conflict
between Conservation and
Exploitation. University Press of Florida, Gainesville.
MAP DATA SOURCES

Map
16.20 Great apes data are based on the following source:
Butynski, T.M. 120011 Africa's great apes.

Stevens,
E.F.,
Arluke,
A.,
In:
Beck, B.B.. Stoinski,
eds. Great Apes
and Humans: The
T.S.,
Ethics of Coexistence.
Norton,
T.L.,
SmAhsonian
B.,
Rowan,
A.,
Institution Press,

With additional data by personal communication from Plumptre,
Kalpers,
Williamson,
J.,
E.A..
A.,
A. 120041
Nzamurambaho,
A., Lola, N., Mugiri, G. 120031 Gorillas in
the crossfire: population dynamics of the Virunga mountain gorillas over the past three decades. Oryx 37: 326-337.
Plumptre,
A.J.,
Series
2.
Cox,
D..
Mugume,
S. 12003)
Wildlife Conservation Society,
The Status of Chimpanzees
New
in
Uganda. Albertine
Rift
Technical Report
York.
ACKNOWLEDGMENTS
Many thanks
to
Maria Arnold lAustralian National University],
Thomas Butynski
IConservation international), Alexander
Harcourt lUniversity of California, Davis], Joanna Lambert lUniversity of Oregon], Julia Lloyd IJane Goodall InstituteUganda), Cherie Montgomery-Lianda IChimpanzee Sanctuary
IWildlife
&
Andrew Plumptre
Wildlife Conservation Trust),
Conservation Society], Vernon Reynolds IBudongo Forest Project], Craig Stanford lUniversity
California],
and Richard Wrangham IHarvard University]
for their valuable
comments on
of
Southern
AUTHOR
Nigel Varty,
UNEP World
UNITED REPUBLIC OF TANZANIA

Jared Bakusaand Edmund McIManus

The United Republic
because
was
Tanzania's neighboring
Tanzania
is
forged from the union
so called
in
1964
of
countries are:
Burundi,
Rwanda, Uganda, Kenya, Mozambique, Malawi,

Zambia, and the Democratic Republic
of the
Congo
Tanganyika and Zanzibar, shortly after their inde-
IDRC].
pendence from UK-administered UN trusteeship
36 million people, with an annual growth rate
in
1961, and from the
Situated
412
it
of
in
UK
in
1963,
respectively.
East Africa, bordering the Indian Ocean,
It
has
1.95 percent.^
population
The land area
of
is
approximately
of
886 037 km^ and
agriculture accounts for half of the US$10.1 billion
AFRICA: United Republic of Tanzania
gross domestic product IGDPI'', or about US$280

per person per
''
year,''
provides 85 percent of
Box
13.2).
This can be explained
exports, and employs 80 percent of the worl< force.
growth rate
Topography and climatic conditions, however,
2.8
limit
cultivated crops to only A percent of the land area.
Tanzania
and
Vi'orld,
the
one
is
in
poorest countries
of the
2004 was ranked 162nd out
Human Development
Index." Gold,
exploitation, however, along with
is
expected
Tanzania
is
177 by
of
and gas
oil,
growth, which
be more than 5.2 percent
to
the
macroeconomic
GDP
reforms, have helped support
in
2004.'
in
peaceful and stable with few tribal or
in
part by
in
the
Kigoma region increased from
percent 11978-1988 census) to 4.8 percent
Gombe NP
11988-2003 census). As a result,
now almost isolated as a forest island,

where once was a small sample of a much larger
(35 km')
is
it
forested
farming
Subsistence
landscape.
marginal areas such as Ugalla
is a
of forest destruction.
In
979,
It
was estimated
individuals,' but
Only the eastern
chimpanzee [Pan troglodytes
schweinfurthii]
naturally
is
present
Tanzania.
in
Chimpanzees generally prefer evergreen or semiwhich
Tanzania occurs most
in
often at lower altitudes and along river and stream

valleys."'
" They are found
and Rukwa
in
the regions of
in
Kigoma
the western part of the country, along
the shores of Lake Tanganyika, and further inland to

the west of the Ugalla River, including the
Tongwe
A small
population of chimpanzees, probably
western subspecies,
duced
nally
caught
d'lvoire
in
and kept
(P.
f.
verus],
Rubondo
was
in
Lake
origi-
Island
Guinea, Sierra Leone, and Cote

in
captivity in
released into the area that
Europe before being
is
now
the
Rubondo
National Park INP) 1457 km'). The population
mated
to
intro-
These 17 animals were
the forested
to
Victoria in 1966-1V69.
some
decline
given the drastic habitat loss
Tanzania, with
in
in
is esti-
have at least doubled since the animals
500-2 500
must be assumed
the areas along the
Tanganyika Lake shore and north
of
Malagarasi
River More than 80 percent of the Luiche, Mlele,
and Mkuti Forest Reserves,
for
example, has been
converted to farmland or burned tor charcoal.'"

Population estimates exist for Ugalla, Lwazi,' and
Gombe
NP,' where
there
are
fewer than
90
and Mahale Mountains NP, where
individuals,"
there were believed to be about 700 individuals
1967." The precise numbers
East Forest Reserve."
of the
were fewer
that there
than 2 000 eastern chimpanzees
estimate was essentially unchanged, at
forest,
in
growing cause
fewer than 480 breeding females.' By 2000, the
regional divisions.'
deciduous
human
population pressure, since the annual population
where
in
of
in
chimpanzees else-
Tanzania are not known, and
it
is
uncertain
whether the population has actually been stable

over this period.
LEGISLATION
The government
of
Tanzania
is
a signatory to the
and the
UN
Convention
(ratified April 1997).
to
(ratified
Combat
1996),
Desertification
Chimpanzees and
their habitats
were introduced."'^''
are legally protected under the National Forestry
THREATS
the Wildlife Policy of Tanzania (1998).
Policy (1994), National Parks Ordinance (1959),
Thanks
to local traditions
opposed
to eating pri-
mates, chimpanzees have historically faced

threat from hunting
in
Tanzania. Recent decades,
however, have seen an influx

countries such as
of
medicine.'
and
DRC
is
''
to obtain
About 388
of
1 1
km'
of forest
in
Gombe
National Park.
kill
remains,
km'
each year,^ but these figures may not

local
deforestation,
environmental
desertification,
and
degradation. Using 1991 and 2003 satellite
images, for example, forest loss outside
NP
Chimpanzees
body parts for traditional
accurately reflect serious
problems
Institute IJGIl
refugees from
declining at an average rate of 910
(0.2 percent)
soil
of
The Jane Goodall
that do have a culture
primate eating. Tanzanians sometimes
chimpanzees
little
and
has been estimated
at 4
Gombe
percent per year" (see
413
World Atlas
Map
Chimpanzee
16.21
rof /
7
\
'
i
f-f
distribution in the United Republic of Tanzania
3Z'E
'-'y
!-
-*
tW^, *Bukoba
Lake
'ft
Musoma
RWANDA
Victoria

Estimated range
end of this country profile
Species
UGANDA
V-
thie
SS-E
i-!
Vi
I,-
Data sources are provided at
^^.j' Eastern chimpanzee
%'
r^
KENYA
^
/X
^
^"
1^
c*)
BURUNDI
^X\
Mwanza
"^^
.^
'"'
r
*
Sereng
NP
^v
<C7 .,,.^
Arusha
Shinyanga
Kilimanjaro
^""-, NP
Mî;
>
UNITED REPUBLIC
OF TANZANIA
/,VCT.I;V
OCE.IX
ZAMBIA
1
MOZAMBIQUE
32E
1>M>
>-
AFRICA: United Republic of Tanzania
Gombe, at the northern end of Lake

was declared a game reserve in 1943
Tanganyika,
and a national park
Gombe NP
in
The chimpanzees
1968.
have been studied continuously for
of
many
years and are well protected." The Jane Goodall

Institute
founded
(JGII,
1977 by Jane Goodall,
in
forest
the
and aquatic
livelihood
biodiversity while strengthening
and
environmental
Mahale Ecosystem Management Project

ing
to
generating
build
activities,
institutions for sustainable
chimpanzees
dialog
Institute
in
the park. To help conserve chimp-
western Tanzania, the Jane Goodall
in
founded the Lake Tanganyika Catchment

Education ITACAREI project,
Reforestation and
which seeks
ation of forests.
to
people
to involve local
It
works with 33
promote the preservation
the restor-
in
villages
and aims
primate habitat,
of
community-centered conservation, education, and
women, and
youth engagement, enhanced roles for

control of the
bushmeat
trade. Activities include the
planting of trees, prevention of soil erosion, and the
promotion
of family
planning and AIDS awareness.
Mahale Mountains
recent assessment of
by the Jane Goodall Institute and USAID. using a

reduction assessment
threat
ITRAI
method and
remote sensing and geographic information system

major direct threats
data, identified five
IGISI
Gombe NR' These were
forests outside
of forests
conversion
subsistence farming, conversion
to
cash crops such as
oil
to
to
palm, local-scale logging,
firewood extraction, and burning. The assessment
concluded that
need
to
more
for
reduce these threats there
strategic
approaches and more

patches. As a
Institute
Design'
is
wider ecosystem, and
and ecotourism.
General management plans for Rubondo,
Mahale, and
ration
Gombe NPs
are currently
in
prepa-
through collaboration between Tanzania
National Parks ITANAPAI, the Frankfurt Zoological

Society,
and the Jane Goodall
Other
field
Rubondo
Institutes.
research programs include one

is
in
currently focused on chim-
distribution,
and another on
Island to look at the ecology of the island,
the social and ecological adaptation of the chim-
panzees
to their
new
habitat,
and their relationship
with other wildlife (indigenous and introduced! on

the island.
Since
Institutes
the
mid-1990s,
the
Jane
Goodall
have helped establish chimpanzee sanc-
tuaries throughout Africa.

Michael Huffman
Kansyana
Valley,
Mahale
National Park, western
conservation-planning
Tanzania. Located along
spatially
is
and temporally
at the scale of specific
result,
now adopting
planning
and
for the
focused conservation actions

forest
priorities
management plans
NP and
panzee ecology and
TACARE project.
the TACARE project
are being developed as part of the
community-based
support park administration, resource protection,
cooking, and therefore alternative sources of energy
in
work-
development, enhance
ecosystem management
threats, develop general
the Ugalla area that
driven
is
is
between the park and the community,
identify
part by a need for fuel for
Habitat loss
of
develop conservation-compatible, income-
provides support for field behavioral research of
anzees
security
park-adjacent communities. To achieve this the
the Jane Goodall

'Conservation
the shores of Lake
Tanganyika, this
is
the
central habitat of the
by
Mahale chimpanzees
process and tools that were
studied since 1965.
the
developed by The Nature Conservancy ITNCI,

order to improve the effectiveness of
its
in
conser-
vation actions.
The Mahale Mountains area was designated

as a national park
his colleagues
chimpanzees since
Society
has
operations
in
in
1985. Toshisada Nishida and
have conducted research on Mahale
been
1965.'^
The Frankfurt Zoological
providing
support
the Mahale Mountains
This support has been
NP
to
park
since 1985.
expanded through the
Mahale Ecosystem Management Project (MEMPI,

funded by the European Union, which
to
is
scheduled
run from 2003 to 2008. The goal of the project
is to
conserve the Mahale ecosystem's outstanding
415
World Atlas
viability of this
Tanzania
not merely a stabilization, of
free of the overwfielnning systematic
is
problems
tfiat
elsewhere
in Africa,
trade,
determine conservation agendas

such as a rampant bushmeat
chimpanzee
habitat
outside the current park boundaries.' Mahale
NP
also faces habitat degradation threats, despite
its
much
widespread and accelerating logging, and
population depends on an increase,
greater size." Both
Gombe and Mahale
warfare with consequent refugee and displace-
chimpanzees also suffer serious threats from
ment problems. Accordingly, conservation needs
disease,
in
Tanzania are mainly
'tactical'
and site-based,
much
which may be introduced by
of
proximity to people and
human
settlements."'
'''^^
revolving around the need to stabilize land use
in
To address these challenges, long-lasting pro-
and around particular protected areas, and
to
grams are needed
expand or restore them as necessary. The chimpanzees
Gombe NP,
of
since the park
is
now
long term,
for
example, are threatened
them
too small to sustain
that
it
surrounded by bare
that should be conceived
discussed with local experts
and should take
areas,
in
into
and
the surrounding
account the dual
the
objectives of arresting the rapid degradation of
has become a forest island
lands and forests, and improving the standard of
hills.''
all
In
in
likelihood, the
living of
the villagers.
FURTHER READING
Goodall,
(19901 Through a Window:
J.
My
Chimpanzees of Gombe. Houghton
Thirty Years with the
l(/littlin
Company,
Boston.
Massawe,
E.T. 119921
Monographs 13
Nishlda,
Assessment
ed. (1990) The
T.,
in
Chimpanzees of the Mahale Mountains.
Tutin, C.E.G., Wfilte, L.J.T.,
fragments
ot the status of
in
western Tanzania. African Studies
35-55.
111:
Mackanga-Missandzou,
University of Tokyo Press, Tokyo.
The use by
A. (1997)
an Equatorial African savanna. Conservation Biology
'\^
rain forest
(5):
mammals
of natural forest
190-1203.
MAP DATA SOURCES

Map
16.21

A.,
Stevens,
E.F.,
Arluke,
A.,
In:
eds, Greaf
following source:
T.S.,
Hutchins, M., Maple, TL., Norton,
B.,
Apes and Humans: The Ethics of Coexistence. Smithsonian
Rowan,
Institution
Press, Washington, DC. pp. 3-56.
With additional data by personal communication from
Idani, G.
and Ogawa, H. (2003), Pintea,
L.
120041,
and from the
following sources:
Moscovlce,
L.R.,
Huffman, M.A. (2002) The chimpanzees
of
Rubondo
Island,
Kakakuona. Tanzanian
Wildlife 27:
56-60.
Ogawa,
H. (1997)
The discovery
Pan Africa Newsi
(1).
Schoeninger, M.J., Moore,
of
chimpanzees
in
the Lwazi River Area, Tanzania: a
new southern
distribution limit.
http://mahale.web. infoseek.co.jp/PANM_lM(1)-01. html. Accessed June 12 2005.
J.,
stable isotope evidence.
Sept, J.M. (1999) Subsistence strategies of two "savanna
American Journal of Primatology ^9
(4):
"
chimpanzee populations: the
297-314.
ACKNOWLEDGMENTS
Many thanks
to
Michael Huffman (Kyoto University),
(University of Oklahoma),
Lilian
Pintea (The Nature Conservancy), Janette Wallis
and Michael Wilson (Gombe Stream Research Center)
draft of this section.
AUTHORS
Jared Bakusa, University
of
Dar Es Salaam
iU
Monitoring Centre
for their valuable
comments on
the
ASIA: Indonesia
Chapter 17
Asia
REPUBLIC OF INDONESIA
Kim McConkey, Julian Caldecoh, and Edmund

The Republic
dence from the Netherlands

Dutch police
territory lies In areas that are not Asiatic in either a
Indonesia declared
of
indepen-
its
1945 and, after
in
was recognized
action', this
by
all in
biogeographic or a cultural sense. Great apes are

restricted to the islands of
Sumatra (475 000 km'l
and Borneo (740 000 km^
which 536 000 km^
1949.
The independence agreennent excluded West
Indonesian
New
Guinea (Papua/Irian Jayal, which remained
which
under Dutch control

sorbed
until
when
1962,
became
unitary
president, Sukarno.
was
dissolved
under
republic
The
Introduction of martial law
this
A coup and counter-coup

suppression of the Communist
the
(PKII,
massacre
of
led
tarian
manner
of the Asian
1965 led
in
from power
crisis of
from the Asian (Sunda) continental shelf
alongside Java (133 000 km'l.
had a
Indonesia
235 million people

rate of
in
the midst
population
total
percent. This low rate reflects long-
1.5
standing government sponsorship of a nationwide
dua cukup
(two's
enough) family-planning strategy.
Nearly two thirds of
all
Indonesians
is
because
civilization,
fertile
of their long history of
hundred
turally diverse, with several

lingulstic
part of
groups - many
what
is
them
of
election,
direct presidential
used
is
were Inaugurated as president and
Kalla
vice
made up
is
of
about 17 000 Islands
with an aggregate land area of nearly 2 million
These islands have some 50 000-80 000

coastline between them. They are set
km^
of territorial
In
Ocean
kml
km
to the Pacific.
of
3 million
sea that extends for 5 100
between mainland Asia and Australia,

Indian
km
linking the
Although traditionally
regarded as a Southeast Asian nation, and indeed a
founder
Asian
member
Nations
of the Association of
lASEAN),
It
for
the
in
formal
arose from
language used
throughout the Malay Archipelago (which includes

Malaysia and the Philippines).
In
area
1988, nearly 10 percent of Indonesian land
was
classified as arable
and
7.2 percent
was
under permanent crops.' Subsistence farming and
president on October 20 2004.
Indonesia
ethno-
West Papua,
culturally the richest island
world. The Indonesian language
Bambang Yudhoyono and Jusuf
Susilo
distinct
in
Malay, an Austronesian trading
first
Bali.
advanced
supported by irrigated farming on their
purposes throughout the country.
Abdurrahman Wahid, and Megawati
on Java
volcanic soils. The Indonesian people are cul-
changing rapidly under the successive presidencies
Sukarnoputri. Following the
live
Madura and
of
then, Indonesian Institutions and society have been
of B.J. Habibie,
about
of
2003' with an annual growth
This
1997-1998. Since
is in
Western Indonesia,
in
to the
an authoriin
Kalimantan)
and on the nearby islands
thousands
of
rise
of
1959-
in
installation of President
US backing
until his fall
economic
the
to
a period of
Party of Indonesia
hundreds
and the
of Its supporters,
Suharto; he ruled with
In
it
1955,
In
communist-Influenced 'guided democracy'

1965.
1950;
founding
held
1957 and
in
In
its
first election,
had an Inconclusive outcome;
was ab-
it
country had a
into Indonesia.' Initially the
federal structure but this
McManus
much
of
Southeast
Indonesia's
fishing
IS
important
to local
people throughout the
many areas have been converted
country, and
agricultural plantations, often of
guinesnsis].
under
oil
oil
palms
Between 1967 and 2000, the
palm plantations
less than 2 000
economy has
km'
to
in
to
{Elaeis
total
area
Indonesia grew from
over 30 000
km"' The
overall
long been dominated by the extractive
industries, however, with
minerals providing
much
oil,
of
gas.
and hard-rock
the nation's foreign
exchange. The timber Industry
is
also important.
417
World Atlas
MAP
17.1
in
Indonesia
this
country profile
PAPIIANEW
GUINEA
^Ci
S g )
2 "5
5
<
J
-
Q<
,'
-1
Xj
i^
Li
3
^
;:
<^
Q
o
ea
<
-h
Q
Z
<
V
E
j
a.
-<
X
H
en
CD
I"
J,''
>
QJ
3
"
o in in II.
B s
= S >" 7 7?'"
2 S o in o in CM
ii -
618
"
<-
*-
III
ASIA: Indonesia
having begun
the early 1960s
in
Kalimantan; by
in
concessions had been awarded covering
1988,
about kZU 000 km^ or three quarters
of the total
Indonesian forest estate.' Follovîng a governnnent

decision
in
the 1970s to phase out exports of logs,
Indonesia exported just
m^ as
million
\.li
logs and
8.2 million m''as plywood in 1991.'The total log har-
meanwhile, rose from about 16 million m^
vest,
1
m^
981 to about 26 million
get harvest for 1995-2000
in
987;' the
annual
was over 37
1997, and the
fall of
in
tar-
million m'.
financial crisis, however, intervened
The Asian
knowledge
the Suharto regime
in
in
1998
of the
population size and rate of change
within each."
Sumatra, 13 habitat blocks have been iden-
In
there were thought to be about 7 000
2003,
In
orangutans inhabiting these areas," with three

populations each containing over
Orangutans appear
to the
south
of
508 individuals]. East Leuser
the Tripa
processing industries
annual allowable
was 63
shortfall
logs annually, resulting
Rupiah 30.42
least
lost taxes,'
in
is
while the
government, was
made up
being
which produces 50.7 million
illegal logging,
at
m^
million
cut, set by the
m\" The
12 million
estimated that
is
It
by
m'' of
state financial losses of
in
trillion
(US$3.18
billion!
in
and putting overwhelming pressure on
settlements along the coast of Aceh,
ands
little
of people.
It
is
affected, but the resultant
for rebuilding
thous-
killing
thought that orangutans were
demand
poses a new threat
northern Sumatra (see Box
timber
for
to the forests of
to
make
priority is to
a single popu-
Swamps
difficult
reconnect to West Leuser, but a degraded forest
corridor
still
exists
Trumon-Singkil
between West Leuser and the
Swamps
that
by the Leuser Development
being rehabilitated
is
Programme (supported
by the European Commission).
West Kalimantan, the 2002 estimate
total available habitat for P.

to
about 85 000
tat
km^"
blocks and
population
of
in
is
for the
pygmaeus amounted
between 306 habi-
divided
inhabited
individuals." There
estimate
11.21.
(280 individuals).
about 5 340 orangutans. The Tripa
lation of
In
tsunami wave destroyed
052 individuals),
(1
500 individuals), and
(1
are already very fragmented and would be
''
December 2004,
Swamps
Swamps
reconnect these four units to
Indonesia's remaining protected forest estate.'

In
two
of
contains four habitat blocks: West Leuser
11.21
(2
regulate the timber industry.
timber from Indonesia's wood-
locally extinct
small populations. The Leuser Ecosystem (Box
the Trumon-Singkil
for
000 individuals.
have become
to
Lake Toba, with the exception
seriously reduced the ability of the government to
2002 the demand
kml
representing together almost 9 000
tified,
by only 2 000-2 500
a relatively large P. p.
wurmbii
Central Kalimantan, with a current
over 32 000. This includes the areas of
Tanjung Puting, Gunung Palung, and Sebangau

Indonesia
is
home
to
National Parks (NPsl,' and the
abetii in the northern part of
Sumatra and
subspecies of Pongo pygmaeus
East Kalimantan, P.
and
also
Map
P. p.
p.
wurmbil
17.1.1 P.
all
three
Kalimantan
in
also occurs
in
Sarawak
habitat,
occurs
in
the
in
of
parallel with the reduction

in
major rivers or wide swaths
use
habitat
is
separated from
all
of cultivation.
habitat
block therefore corresponds to a separate popunot easily colonized by individuals
from other populations. Once habitat blocks are

conservation
total
Schwaner Mountains."'"
wurmbii. The main population of
Kalimantan
Gajah),
is in
P. p.
morio
the Berau area (including
where an estimated
approximately
p.
East
in
Gunung
558 orangutans
survive."" Several smaller populations

ing
It
area of 5 600 km^ of dipterocarp forest,
500 individuals
to
exist,
add-
this
sub-
species population.
Indonesia.
conservationists
others by normally impassable barriers, such as
identified,
area, sur-
Arut-Belantikan
with an estimated population of 6 000-6 500 P.
area of connected habitat that
is
is
West Kaliman-
blocks' as their basic planning unit, each being an
one that
2003,
in
Central Kalimantan. (See
in
the distribution of forest
lation,
the foothills of the
in
pygmaeus
and most suitable
Orangutan
time
has a
Indonesian sector of the island.^" The distribution
in
in
first
the province of
pygmaeus
the apes has declined
veyed for the
Kapuas-Barito
An important new
in
(Malaysian Borneo), but 80 percent of the population,
floodplain (Mawas).^'
mono
(Indonesian Borneo): P. p.
tan,
Pongo
both orangutan species:
planning then
requires
THREATS
Consistent with the nature of the Indonesian land-
development process, which has been based largely

on logging and the expansion
of plantations,
orangutan species are threatened mainly by
both
signifi-
cant and ongoing habitat loss and forest fragmentation,^"
aggravated by hunting and persecution as
agricultural pests."
These continue, despite the
that orangutans have
been protected
in
fact
Indonesia
419
World Atlas
since 192i. Conservation efforts fiave failed to slow
decline of orangutan
tfie
tiabitat.
About 55 000 knr
breeding habitat for Bornean orangutans was
of
between 1993 and 2002, and much
lost
remaining forest
fires."
The
logging,
the
of
affected by logging and forest
is
probability of forest fires
areas based on two main categories: sanctuary
which opens and fragments the forest and
The same law further allows

zoning (Articles 32 and
become more prone
dangered and rare species
2010
000
killed
destroyed a tourist village

of
forest
year
2003, a landslide caused a flash flood
Gunung Leuser NP which

regency
little
to survive past the
Sumatra or Kalimantan. '^
either
in
November
Very
ignition.
to
expected
is
Langkat. Other floods
been linked
KO
over
in
of the
Management
wildlife protection are the responsibility
the
development also threatens
government investment
building lobby
IS
in
development. The road-
powerful, conservation interests
are relatively weak, and
numerous protected areas
throughout the country have had
roads
through them. That this danger persists

the Ladia Galaska road project
will,
if
system.
It
and
is
built
shown by
Sumatra, which
in
Leuser Eco-
the
would also open up a very large new area
commercial
for
in
is
exploitation,
whether
legal or not,
hence backed by powerful lobbies within
local
government, business, and the armed forces. As
General
of
Forest Protection and Nature Conservation (PHKA],

in
with
collaboration
ment
The legal and

of
conservation law has never been com-
pletely clear
many
government and the
local
institutional basis for enforce-
in
Indonesia, however, and there are
presidential, ministerial,
ment decisions Ikeputusan]

example,
law. For
made
it
wild orangutans
Bohorok
of
in
1995,
illegal to release
still
and local govern-
that have the force of
Law UU 280/kpts
orangutans
in
11/1995
areas where
persist (leading to the closure
Sumatra as
in
rehabilitation
and
release center].
elsewhere, roads bring with them greater access
who may hunt, and for settlers who

forest and may kill orangutans that raid their
Protected areas
In
Indonesia,
conservation planning went hand-
in-hand with forest and land-use planning. The
development
consensus
timber industry was based on
of the
classification
in
clear
government agencies
newly established crops. Kalimantan's peat-swamp
agreed allocations
ries of
float
10.31 built
out illegal logs, which are draining and
function
forest
1970-1985, based on discussions
forests also suffer from canals (see
Box
of
[Tata
Guna Hutan Kesepakatan, TGHK!. The TGHK arose
for people
to
conservation area
continued, bisect one of the largest remain-
orangutan populations,
ing
of
of the
of the Ministry of Forestry's Directorate
police.
most tangible expressions
20-25], and
areas (Articles
16 and 29].
orangutan habitat, as road building has long been
seen as one
(Articles
for protected
system and
to illegal logging.
Infrastructural
zones
of
en-
in
the region have
in
(Article 18], protection of
In
people and
Lawang
at Bukit
refers to buffer
management
for
the constitution
34!,
Biosphere Reserves
nature re-
grand forest parks, and natural recreation parks!.
allows normally moist forest areas to dry out and
below
strict
vation areas (Article 29, comprising national parks,
increased by
is
U, comprising
reserves (Article
serves and wildlife sanctuaries! and nature conser-
and
use.'
among
various
produce maps showing
of forest
permanent
their forestry
to
land to various catego-
The
five
main categories
uses were:
killing the forests.
The net
result
is
that both orangutan species
are seriously endangered
in
Indonesia, making
urgently necessary to identify locations

is
a realistic
and
chance
it
where there
of protecting viable populations,
to direct appropriate
investments
to
them,
nature reserve (no timber extraction];

protection forest (no timber extraction];
limited
production forest
regular production forest

tion or clear felling,

Consen/ation
in
Indonesia
based on Act No. 5 of
non-industrial
(for industrial
selec-
according to forest type!;
and
conversion forest
is
(for
selection felling!;
(for
clear felling and con-
version to other uses!.
1990, Concerning Conservation of Living Resources
and
their Ecosystems. This lists species (including
This classification system took
little
orangutans! covered by the Convention on Interna-
traditional land claims by local
communities, who
tional Trade in
Endangered Species
of
and Flora (CITES! and defines a variety
iiO
Wild Fauna
of
protected
notice of any
were not consulted.
The categorization program was also flawed
ASIA: Indonesia
due
insufficient information to support spatial
to
and forestry planning,' but
were undertaken
tions
comprehensive
Transmigration
was
tfiese
of
mapping opera-
otfier
correct
to
tfiis.
the late 1980s: this
in
most
Tfie
by the Ministry of
mapped
land
use and land capability for the whole country

outside Java and Bali, with a view to finding suitable
places to receive
The Ministry
sponsored settlers."
officially
Works
Public
of
Transmigration (RePPProTI
the
TGHK
integrated
later
Program
these Regional Physical Planning
maps
for
as well as district and provincial planning
Jambi province,
new remote imagery
vations and
to
field
obser-
show
actual
forest cover, protected forests, nature conservation
sanctuary reserves,
development
alignments
the
as well as proposed
existing
of
and other
roads,
projects. Act No. 2k of 1992 provided
Sumatra.
per
km
per year for the reserve system as a
whole. '^ This can be compared with the 300-500

percent higher levels
nature reserves
in
of
expenditure on priority
Thailand and China, as well
a comprehensive legislative context for a national
as with the recommended
system
US$300-400 per km' per year
spatial
of
although cases
planning,
between planned and actual uses
conflict
of
of land
Planning for a national system of protected
was done in parallel with TGHK,

RePPProT By 1990, Indonesia had
Indonesia
in
and later with
of
gazetted 303 terrestrial nature reserves of various

kinds totaling 160 000 km^ or 8.2 percent of land
area, and another 20 000
km'
at
175 sites had been
to
million per year
in
same general range as
in
Indonesia
the
into
Thailand and China.
The Asian
fall of
the Suharto regime
30 years
ability of central
most ecosystems, and populations
most
native
TGHK and RePPProT
been designated through
The main
was
as
more than
power, caused grave disruption
in
for forest
government
to
management
to the
impose an orderly
or conservation on
the outer islands, such as Sumatra and Kalimantan.
Indonesia
is
organized into provinces, kabupaten
regent or resident); kecamatan (subdistricts, each
slopes.
Conservation efforts
action
plan' and
analyses such
policy
headed by a camat]; and various kinds

munity As
consequence
of the political
of
com-
events of
the late 1990s, a significant transfer of authority
Indonesian Country Study on Biotogicat
Diversity.^'
donors
The
1998, after
the safeguarding of both water catch-
during the 1990s were guided by a national biodiversity
that
(regencies or districts, each headed by a bupati,
ments and steep
as the
system
in
were
role of these designa-
protection forest.
tions
of
More than 300 000 km^ had meanwhile
meant
derailed.
financial crisis in 1997
efforts to increase investment
proposed as such reserves.'" These areas were sel-
species.''
would have been required
have brought average expenditure rates
ected to include viable and representative samples

of
minimum figure of
for the management
national parks, suggested by lUCN." At least
US$130
continued to occur
areas
in transit
to their release site in
maps. These were then updated using
areas,
Orangutans
with those from
These were being supported by
year The development
for
an effective system
of
protected areas therefore
good coverage
official
about US$12 million per
to the extent of
seemed
of
possible, giving
most components
of
Indonesia's
A young orangutan
at the
Sumatran
Programme's
quarantine
site.
biodiversity.
Many reserves had
little
effective
manage-
ment, however, reflecting the limited resources

the
responsible
of Forestry, the Directorate
tection
General
of
Forest Pro-
and Nature Conservation (then the PHPA,
now the PHKAl. At

PHPA and its
the
of
department within the Ministry
that time, rates of expenditure by
partners averaged about
US$75
621
World Atlas
Heritage
of
Gunung
Site.'^
NP
Leuser
is
Leuser Ecosystem area and
the
NP
orangutans, while Kennci-Seblat
is
part
supports
the site of
reported 'orang pendek' sightings Isee Chapter
1).
Both national and international nongovernmental
organizations
(NGOsI found
hard to work
it
in
Indonesia under the Suharto regime, as they tended

to
be viewed with suspicion by the hard-line mili-
tarists
who held most power at that time. They

some political protection, however, from
received
the Ministry of State for Population and Environ-
ment IKLHI and
Some became
long-time Minister Emit Salim.
its
with central government
Bornean orangutans
(e.g.
WWF-lndonesia
the
program], by gaining the strong support of pro-
at
governments
the Wanariset
vincial
Rehabilitation Centre,
Birute Galdikas
in
Rondang Siregar
and
research,
from the center
to the regions
considerable authority granted
make
use.
in
own
their
The
to
was agreed,
with
the kabupatens to
decisions regarding land and forest
result in
many areas was
a rapid increase
both
at
by
Conservation,
levels.
relating
activities
to
sanctuary
common
rehabilitation or reintroduction often had
roots during this period, although they have since
become somewhat
differentiated.
They can now be
classified roughly into:
logging and forest clearance, both legal and
quarantine
illegal."
The kabupaten
enough
to
its
inhabitants, but small
gramme
however, that local people are becoming more en-
public
new
response
utan Survival Foundation (BOSFj;
and reintroduction
rehabilitation
demand, some kabupatens have proposed
such as those
at
in
Lemandau
in
Kalimantan; as well as
the most recent example, and
Tiga Puluh
NP
in
to
protected areas to Jakarta. Batang Gadis
reflects
is
NP
long-suppressed wishes, expressed as
field
research on wild orangutans,
Cabang
Leuser Ecosystem. Similarly,
Kutai, Tuanan, or
local people
East
in
the
Global Conservation
habitat-conservation
Organization.
under control.
In
2004,
logging
jail
or the
sentence
jects
it
with
12 years, or by death
in
starting
of
of
fires
In
July 2004, an area of 25 950
km'
jointly
covered by the Gunung Leuser, Kerinci-Seblat, and

Bukit Barisan Selatan
Tropical Rainforest
NPs was designated as
the
Heritage of Sumatra World
Kalimantan; and
in
projects,
such as the
at
Mawas, Tanjung Puting NP, and
Gunung Palung NP.
In
Sumatra, the
program
exceptional cases.
in
for
Leuser Development Programme, and pro-
proposed a law that would punish convictions for
minimum
Mawas
at,
Puting,
Sumatra]; and
Meanwhile, central government has been attempting to bring illegal logging
Tanjung
Panti.
Ketambe, Bohorok, and Dolok Sibual-Buali
Kalimantan had
lobbied for a national park for decades, supported
WWF-The
at Bukit
Jambi province, Sumatra;
in
example,
in
activities,
Meratus, Tanjung Puling, and
early as 1928, by traditional leaders to protect the
Kayan Mentarang area
of
(SOCP], the Orangutan Foundation
local
northern Sumatra
illegal
such as those
for,
International and UK, and the Borneo Orang-
connected individuals. There are signs,
vironmentally assertive, and
which formerly captive
Sumatran Orangutan Conservation Pro-
the
be influenced by private companies and
rich or well
activities, in
orangutans are cared
a political unit too large to
is
be easily accountable to
by
run
the operation
(e.g.
Central Kalimantan], or through
making alliances
Indonesia.
422
by working carefully
established
It
in
SOCP
has established a release
Bukit Tiga Puluh
NP
in
Jambi province.
also conducts most of the survey and monitoring
work concerning the status and
distribution of wild
Sumatran orangutans. The Research, Monitoring,

and Information Division
of the
Leuser Management
ASIA: Indonesia
Unit
manages research
activities within the
Leuser
Ecosystem.
International and
UK
fund patrols
Foundation
Tanjung Puting
in
NP, rehabilitate and release orphan orangutans
in
Lamandau Nature Reserve, and support research

Into conservation and forest restoration. The BOSF
and releases orphans
rehabilitates
papan area and

Involved
5
in
other parts
peat-swamp
of
Balik-
Kalimantan, and
of
Mawas
Is
area,
forest inhabited by orang-
NGOs
Several partner
utans.
the
In
proposals to protect the
in
000 km'
other countries
In
include Balikpapan Orangutan Society-USA (BOS-
who
USAI,
provide support for the
work
BOSF.
of the
The Orangutan Conservation Forum (OCFl

group
tions with educational programs.

ginally
is
orangutan-focused conservation organiza-
of
need
efforts
Kalimantan, the Orangutan
In
protocols are standardized. Collaborative field
planned at a meeting
Central Kalimantan,
In
2002;
The OCF was
Palangkaraya,
In
It
ori-
alms
as a
to act
centralized body for communication and facilitation

of the sharing of Information
and Individuals Involved
between
all
as well as other environmental education.

also play a key role
In
groups
orangutan conservation,
in
It
will
advising upon the Indonesian
to
be
at
be
Ketambe
particular, research
In
the Leuser Ecosystem should
in
and expanded.
continued
Forest
loss
should be monitored and a sustainable source

of
funding for long-term orangutan research
in situ
should be sought. Participation by local
NGOs should be encouraged.

Protection.
needed
training
techniques
the
In
and protect populations,
identify
to
and enforcement
Surveillance
teams require
keystone resources, corridors, and essential

outside current protected
habitats
areas.
Helicopter surveillance should be considered.
Conservation policy should be integrated Into
governmental policy
Education. The development of an Increased
awareness
via
of preservation
education programs
national
In
needs
required
Is
both schools and
media
International
Institutions.
coverage of the current status
orangutan
of
populations should be sought.
Regional actions
Sumatra. Restore the
in
connections between
national great ape survival plan INGASPI.
with umbrella pro-
Initiated,
jects for both islands.
blocks
habitat
the
in
Leuser Ecosystem and surrounding areas
(i.e.
connect West and East Leuser, and Trumon-
The forces pressing towards the complete destruc-
Slngkil and
Sumatra and Kalimantan
tion of lowland forests In
are very powerful, and the outlook for species that

inhabit those forests
of hope, in that
to
demand
Is
people
worrying. There are signs
some areas
In
protected areas and local government
leaders are starting to grant
dorsement
ness
in
are starting
them
with the en-
of the national authorities. Public
of the plight of
Indonesia, and
is
willingness to act
against illegal logging
political
Is
rapidly Increasing
becoming established. The
urgency for action varies
among
depends upon the current
rate of logging
the orangutan population. For

the need for action
to survive.
If
Is
habitat units and
some
Immediate
If
size of
habitat loss can be controlled, then
relevant and valuable.
action
and
habitat units,
orangutans are
actions to reduce fragmentation will
for
aware-
orangutans
Some
presented
in
of the
the
become more
major
2004
priorities
Orangutan
West Leuser habitat
and
blocks],
ensure that the Ladia Galaska road scheme
does not pass through Gunung Leuser NP.

Regional actions
Puting
NP
of the
to include the north
Sekoyner River and

the eastern forest.
been cut
Fill in
Regional
of the
the canals that have
swamps,
peat
Into
Tanjung
shore
to establish a corridor to
Illegally cut logs in the
Establish
Kalimantan.
Central
in
Extend the northern boundary
actions
out
float
to
Sebangau catchment.
East
in
nature reserves
at
Kalimantan.
Sangkulirang-
Mangkalihat and Sebuku-Sembakung.

Regional actions
capacity at
encouraged
in
West Kalimantan.
Build
Gunung Palung NP. The BOSF
ing Innovative
is
continue developing and refin-
to
models
for the sustainable
servation of the
Mawas orangutan
Development.
Improve
habitat
con-
population.
quality
in
Population and Habitat Viability Assessment Final
degraded areas, possibly through enrichment
Report" are
planting. Increase sustainable
listed below.
natives for
Studies. Systematic surveys throughout the
orangutan range are needed

priority
sites
are
to
identified
ensure that
all
and research
economic
communities surrounding
alter-
critical
orangutan habitats.
Rehabilitation
should
be
and translocation.
given
to
conserving
Priority
the
wild
423
World Atlas
population (efforts In
as opposed to ex-
situ],
care (conservation
captive
Retiabilitation centers
ex
efforts
situ].
should be licensed and
made
proved and
International
policy.
Species concept
freely available.
An
Coordination.
committed by NGOs
Palangl<araya to
Orangutan
ments
Scientific
Vi/as
at
initiate
Indonesia,
of
The World
Forum
utans as one
of
Heritage
endorsed; the governMalaysia,
Saraw/ak are encouraged
Commission should be established, and the

plans for the Orangutan Conservation
in
this process.
Record keeping should be im-
monitored.
realized using funds
the 2002 meeting
the worlds
Sabah, and
promote orang-
to
first
such species.
FURTHER READING
Curran, L.M., Trigg,
forest loss
Jepson,
in
McDonald,
S..
A., Astiani, D.,
Hardiono,
Siregar,
Y.,
P.,
Caniago,
I.,
Kasischke,
E. (20041
Lowland
protected areas of Indonesian Borneo. Science 303. 1000-1003.
MacKinnon,
Monk, K.A. (2001) The end
of
Indonesia's low/land forests. Science 292: 859-861.
P.,
Jarvie,
J.,
Robertson,
J. M.Y.,
van Schalk, C.P. (20011 Causal factors underlying the dramatic decline
K.,
of the
Sumatran orang-utan.
Oo'x 35: 26-38.
UNESCO
Tropical
(20041
31 &id_site=l 167.
Rainforest Heritage of Sumatra. World
van Nieuwstadt, M.G.L., Shell,

of
Heritage.
http;//vi;hc.unesco.org/pg.cfm'f'cid=
Accessed December 5 2004.

D.,
Kartawinata, K. (20011 The ecological consequences of logging
East Kalimantan, Indonesia. Conservation Biology ^5
(41:
in
the burned forests
1183-1186.
Also see Global Forest Watch resources on Indonesian deforestation: http://wvvw.glob.3lforestw/atch.org/english/
indonesia/maps.htm.
MAP DATA SOURCES

Map
17.1
Orangutan data are based on the following sources, with additional information by personal communication
from Meijaard,
E.
12005]
and Singleton,
Riswan (2004) Orangutan
Dadi, R.A.,
(20051:
I.
distribution polygons: developed at the
Leuser Management Unit as part
Leuser Development Programme, funded by the European Commission and the government
Leuser Management
Unit,
Priatna, D., Rijksen, H., Riswan, Robertson,

J.,
I.
Main sources of
Additional information from Dadi,
R., Griffiths, M.,
Sumatra, Indonesia. Based on technical
van Schalk, C, Idrusman, Singleton,
field data:
Bloxam, C, Kuswandono, Long,
B.,
Y.,
I.,
Wich,
S.
criteria set by Singleton,
McPherson,
J.I,
of the
of Indonesia.
Bogor Expedition
and members
of
the Leuser
to
Sumatra (Burton,
Management
Unit's
Antipoaching Unit.
Meijaard,
E.,
Dennis,
Meijaard
R.,
& Dennis
Singleton,
12003)
I.
(2004)
Borneo Orangutan PHVA Habitat
and amended by delegates
at the
Units:
Composite dataset developed by
Orangutan PHVA Workshop, Jakarta, January 15-18
2004.
Singleton,
I.,
Wich,
Byers,
S.,
Husson, S Stephens,
eds 12004)
0.,
S.,
Utami Atmoko.
S.,
Leighton,
Orangutan Population and Habitat
M Rosen,
Viability
N.,
Traylor-Holzer,
K.,
Lacy,
R.,

Please also see
full
acknowledgments
for the
Sumatra data
in
Chapter
11.
ACKNOWLEDGMENTS
Many thanks
Programme)
to
Ashley Leiman (Orangutan Foundation) and Ian Singleton (Sumatran Orangutan Consen/ation
for their valuable
comments on
the draft of this section, and to Mike Griffiths iLeuser International
Foundation) for information on the Leuser Ecosystem.
AUTHORS
Edmund McManus, UNEP World Conservation Monitoring Centre
Kim McConkey,
Julian Caldecott,
626
ASIA: Malaysia
MALAYSIA
McManus
Kim McConkey, Julian Caldecott, and Edmund

Malaysia comprises most of
Sarawak
tlie
Malay Peninsula
(West Malaysia] and the norttiern and norttiwestern
Borneo. Of
parts of Borneo lEast Malaysia), with the southern
fied
of
from the
UK
in
parks, wildlife sanctuaries, and
his successors,
come
and was
be
to
of
a colony
only briefly after the Second World War),

in
North
and Singapore
1963),
(which then seceded from Malaysia
1965).
in
The
federal constitution allocates roles, rights, and responsibilities
among
ments and,
in
the state and federal govern-
particular, gives the states control
posed
and forest management, therefore, are primarily the
Malaysia
nomy
of
is a
governments.
began
peat-swamp
the
in
1970s
plain; in the
interior which,
dominate
late
by the late
diversified into high-technology
1970s
to
and gas. The country's
about 328 550
km^ and
its
total land
population
area
is
was about 23
Sabah
and about
much
managed
in
ways
and
agrarian or else under large plantations (mainly of
and laws
Taman Negara
Krau Wildlife
National Park (NP), and
Reserve).
Although
relatively sparsely populated.
areas
(e.g.
it
Miri in
shifting cultivation
Dayak peoples
(e.g.
Sarawak)
is
has urban
Kota Kinabalu and Sandakan
and Kuching, Sibu, and

under
Malaysia
East
it
Kadazan-Dusun
in
oil
in
palm and other
of
war
in Iraq.
by the growth
valuable
to forest
bio-
management. This process was matched

nature-oriented tourism within
in
the realization that wildlife -
if
used
to
in
-and forests could prove very

new and nondestructive ways.

The
Bornean
northwest
for
pygmaeus pygmaeus] occurs
important
northeast Bornean orangutan
by the sudden
acute respiratory syndrome (SARS) epidemic and by

the
and
diversity
Sabah, Iban,
economy grew by '1.9

difficult first half when
was shaken
Sabahan
more sustainable approach
of a
especially orangutans
protected areas. Malaysia's

2003, despite a
awareness,
Sabah's bio-
largely
timber production, plus a number
in
now
growing env-
to
development
is
managed
investor confidence
severe depletion
developed policies, plans,
promote the survival
Sabah, leading
plantations, or natural forests mostly
percent
From the
of
to
Sabah
and Bidayuh, and several 'Orang Ulu' peoples

Sarawak), or large, recent,
Sabah was
about I960;
logical resources, contributing to the
in
managed by indigenous
in
response
institutions have steadily
Mountains,
until
of this is logged.
that resulted
ironmental concern
the
of
remains under
half
timber reserves,'" and they are
series of large towns, cities, and industrial zones,
Range
km'
1960s into the 1990s, Sabah's forests were
while the interior and eastern parts remain largely
Mam
the
in
the late
only 25 percent of land area remains under
virtually exhausted.' In
the
in
palm and pulpwood
plantations continue to expand. Eastern
year' The western part of the peninsula contains a
(e.g.
oil
almost completely uninhabited
late
to
(73 371 km') occupies 10 percent of the
island of Borneo,'
of the state's
palm) or natural forest
per year
5 million m''
million in 2003, growing at nearly 1.9 percent per
oil
into the hilly
had come
Sarawak has been logged each yean
lowland forest, and
oil,
move
1980s. Since the early 1990s, over 2 000

forest in
now
forestry,
to
1980s,
K million m' per year
about
manufacturing, tourism, and other services, from a

plantation agriculture, production
logging
log production. Total production increased
from about
steadily
major
when
forests of the coastal
began
this
strong base
in
Sarawak has been
exporter of timber since the 1960s,
natural forests,' although
middle-income country, the eco-
which has
for protection.'
nature
been pro-
reserves], while another 6 percent has
over most aspects of land and forest use. Wildlife
responsibility of the state
is
(national
Malaysia!. Sarawal< (which had
Borneo (renamed Sabah
as permanent forest estate, 3.3 percent
the former British
been independent since the 1840s under the Raj
James Brooke and
of
classi-
is
under state law as protected areas
1957, and has increasingly
known as Peninsular
largest
island
37 percent
total land area,
its
the
of
gazetted
It
become independent
colonies of Malaya (which had
Malaysia's
is
in
South China Sea between them.

1963 through a federation
was formed
500 km']
(12
and occupies 17 percent
state
in
Sabah. (See also

In
Map
Sarawak, and the
[P. p.
morio] occurs
17.2.)
Sarawak, significant populations
utans occur only
in
[Pongo
orangutan
in
of
orang-
the south-central interior,
and around the Lanjak-Entimau
in
Wildlife Sanctuary
on the border with Indonesian West Kalimantan
425
'
World Atlas
MAP
17.2 Orangutan distribution in Malaysia
this
country profile
J
a.
en
> 2
c S
=5
ra
c
c S
1 ^
Q.
<D
o
"^
in
1-
in
CN
in
in
=5
3. D-
^
^
o
lU
u
z
-ij
o
ss
to
<
O;^'":
my
626
<-
in
*^
III
ASIA: Malaysia
(which has a contiguous population
and possibly
(over 30 animals),
forest
in
pockets
in
swamp
of
mouths
the coastal region and
of
the
Lupar and Saribas Rivers. Wandering individuals

are
occasionally encountered
the
Tamabu Range (Pulong
Darussalam. The locations
as
as far north
and
Taul
of the
Brunei
source popu-
these wanderers and therefore tor the
lations, for
subspecies
[P.
pygmaeus
p.
More than 60 percent
the Betung
in
Kenhun NPI.'' Small orangutan populations have

been documented at Sedilu (two animals], Sebuyau
mono],
or P. p.
in
live
Sabahs orangutans
of
outside protected areas,
production forests
in
been through several rounds
that have
extraction and that are
still
timber
of
exploited for timber'
The two largest orangutan populations are found

logged production forests
in
in
the eastern parts of the
Sabah Foundation (Yayasan Sabah)
con-
forest
cession [with 3 300-11 900 individuals) and on the
north
side
the
of
Upper Kinabatangan
(approximately 2 300 orangutans,
000
the Deramakot Forest Reserve).'
of
River
them
in
addition, the
In
northern interior Sarawal< and Brunei Darussalam
Trus Madi, Ulu Sungai Milian, and Sapulut forests
are not known. The virtual absence of orangutans
together host perhaps 520 orangutans; and the Ulu
may
north of the Rajang River
be a consequence of
prehistoric hunting or of insufficient fruit supplies
or a combination
populations,
Certainly,
orangutan bones are
common
the de-
in
northern Sarawak, indicating a hunting history
spanning several thousand years. Modern hunting,
settled
extensively
southern Sarawak, has greatly reduced
much
orangutan populations;
been converted
with
who have
people
by Iban
especially
little
mosaic
to a
of this area
Silabukan Forest Reserves together have about 500
orangutans
in total.
Orangutan abundance
correlated
directly
production forests
in
type
the
with
management. The highest orangutan

were
identified
is
forest
of
densities
those forests implementing
in
reduced-impact logging, showing that uncontrolled

logging
have
activities
negative
impact on
orangutan abundance.
has also
of shifting cultivation,
many
intact forest in
Tungud, Lingkabau, Bonggaya, Ulu Kalumpang, and
both factors.
of
Caves National Historic Monument
posits at Niah
in
breeding
to sustain
these dipterocarp-rich forests
in
places. Elsewhere,
THREATS
Hunting
Malaysian Borneo
in
largely of
is
only
logging has also had a negative effect on orangutan
historical (or prehistorical) significance in helping to
population numbers.
explain the
in
Sabah, orangutans are patchily distributed
over their range. The greatest abundance of orangutan nests
500
is
recorded
above sea
lowland forests below
in
Recent aerial and ground
level.
censuses estimated that about

(95
percent confidence
were present
lations with
in
11
000 orangutans
8 000-18 0001
intervals:
16 major populations
more than 50
popu-
(i.e.
degree
individuals, their
of true isolation
from one another being largely
unknown), mainly
in
the eastern and central parts of
the northern and western parts of Sabah,
only two significant (though small and
populations are
now
found:
in
Park and World Heritage
Site,
isolated)
Crocker Range
NP, with about 180 orangutans, and
in
Kinabalu
with about 50. The
Sepilok Orangutan Rehabilitation Centre
Sabah also has about 150 ex-caotive
in
eastern
rehabilitants.
The main protected populations include those

in
the Tabin Wildlife
individuals).
tuary
(1
all
modern
Reserve (with about
Lower Kinabatangan
Wildlife
285
Sanc-
125 individuals),^ Kulamba Wildlife Reserve
(730 individuals), and
Area [730 individuals).
Danum
Valley Conservation
distribution of orangutans, as
the surviving
Sarawak population
is
now
within secure protected areas, and there appears
never
have been
to
much
hunting
in
central and
eastern Sabah. Other, very serious, threats are at
work
in
Sabah; these are interactive, cumulative,
and potentially devastating

factor
to
orangutans. One
the conversion of forest to plantations,
is
particularly of
mangium;
palms [Eiaeis guineensis] and
oil
pulpwood trees such as Acacia
fast-growing
this
has occurred or
enormous areas
Sabah.'
in
almost
of fertile
is
soil in
proposed over
eastern Sabah.
Forest conversion has contributed to the loss of at

least 35 percent of
1980s.''
'An
orangutan habitat since the mid-
equally
severe
threat
is
logging,
especially multiple re-entry logging of dipterocarprich forests.
third
severe threat
is fire, to
which
both logged forests and pulpwood plantations are
dangerously
in
Sabahs seasonal
when droughts are prolonged
vulnerable
climate, especially
during El Nino climatic events.
The current network
of
protected areas
in
Sabah harbors about A 500 orangutans, representing about
3'i
percent of the total
number found
in
the state.' These populations are very fragmented;
427
World Atlas
Sanctuary 1260 km^l, which
adjacent to 257 km^ of
is
unprotected and managed forest; and the Kulamba
Reserve
Wildlife
Orangutan
established
in a
Sanctuary (see Box
Danum
Valley
Danum
the long term, nnost of ttiese are vulnerable to
through
extinction
drought,
the
effects
is
desig-
officially
protection forest reserve."
Valley orangutan population
The
connected
is
Sabah Foundation
concession production forests.
inbreeding,
of
disease, or localized hunting.'
fire,
(438 km^)
protected area within the Sabah
with those inhabiting the broader

in
the
Finally,
10.2).
Conservation Area
nated as a class
Kinabatangan.
the
in
and around the Lower Kinabatangan
in
Foundation timber concession and
Ficustree,
was
(KOCP)
eastern Sabah, esp-
of
Wildlife
set aside as a
A Bornean orangutan
Kinabatangan
Project
1998 to secure the population
in
Kinabatangan floodplain
ecially
The
1204 km').
Conservation
Other protected areas containing orangutans
Range NP
are the Crocker
(2
400
km-'),
Kinabalu
Park (750 km'), and the Sepilok Orangutan Rehab-
LEGISLATION
Most
Orangutans are protected
under the Protection
Sabah
by the
11997),
and
in
Peninsular Malaysia
Enactment
Conservation
Sarawak by the Wild
in
Sabah
of Wildlife Act (19721, in
Wildlife
Ordinance 119981. Their habitats
Life Protection
may
also be pro-
various ways under forest law.
tected
in
parl<s
are gazetted under the Parks Enactment
11984);
Centre (43 km').
ilitation
Legislation
Sabah,
sanctuaries are gazetted
wildlife
Conservation
the Wildlife
In
Sarawak, parks,
wildlife
Enactment
sanctuaries,
under
(19971.
In
or nature
official
Reserves Ordinance
in
in
of
various production forest landscapes
in
Sabah Foundation concession
(an
area of over
4 000 km');
the
to
the
Silabukan
Some
populations
that are sig-
ment
of
areas,
where
(250 km');
1
between them, these
are only minor problems
become serious
illegal logging is
if
in
these
rampant.
8 percent of Sabah's land area
is
reserves.'
the 500-1 500 range, and are there-
fore particularly significant for the conservation of
orangutans.''
(1225 km');
'*
These are the Tabin
the
Centre
in
Kubah NP, the
Rehabilitation
Centre
in
Centre
in
Sabah are
all
involved
rehabilitation of confiscated
in
the care and
young orangutans.
Several of the
areas protected under state law contain orangutan
428
Sarawak, and the Sepilok Orangutan Rehabilitation
the system of national parks and other
in
move-
habitat has been recently destroyed."
Wildlife
categories of existing
to sustain
a result of the
iOG orangutans. Hunting
Wildlife
where
is
refugee orangutans from surrounding
Semenggoh
to
forests (Tabin,
be too dense for the habitat
especially because the areas are contiguous with
populations
managed
Sanctuary (1870 km'), and the
in
The Matang
in
in
of increasing in future. Others,
not monitored,
Close
the Ulu
Trus Madi, and Sabah Foundation) are thought
to
illegal logging
included
in
Reserves (covering 2 621 km'
Forest
the long term. This
areas, but could
Indonesia,
and
in total).
them
contain an estimated
and
total);
in
thought
NP
(Ulu)
the Trus
Tungud, Lingkabau, Bonggaya, Ulu Kalumpang, and
orangutan conservation: the Lanjak-
Wildlife
Ai
in
however (Kulamba and Lower Kinabatangan), are
Sarawak has two protected areas
Batang
Upper
currently to be below the habitat's carrying capacity,
(1998).
Protected areas
Entimau
of
Madi, Ulu Sungai Milian, and Sapulut forests (which
cover about 3 300 km'
and hence capable
nificant for
north
the eastern
Kinabatangan River (some 2 000 km');
reserves are gazetted under the Wild Life Protection
Ordinance (1998) or the National Parks and Nature
Sabah's orangutans do not occur
protected areas. They are instead found
Wildlife
Lower Kinabatangan

The
Sabah
government
recommendations
of
has
recognized
the
an International Workshop on
Reserve
Wildlife
including those outlined below."
in
Sabah,
in
August 2003,
ASIA: Malaysia
management. Sabah's
Forest
forests should
areas, while providing opportunities for the
be managed for orangutan conservation, by
sustainable use of natural resources.
reviewing current and future plans for forest
Protection. Training should
management
sur^/eillance
light
in
strategy formulated
Department,
among
vi/ildlife
by the
Sabah
Wildlife
enhancing
by
relevant
state
of
management
Vi/hich
and
Education. Raise awareness of preservation
management units
harbor over 60 percent of Sabahs
programs
by education
needs,
the forest
in
corridors,
areas should be identified and protected.
by issuing practical guidelines to foresters,

especially
resources,
essential habitats outside current protected
and
authorities,
keystone
lations,
collaboration
be provided for
and enforcement teams. Popu-
media contacts, and through
schools,
in
public
and
governmental lectures.
orangutans.
Agriculture. Agricultural
should
practices
Development. Improve habitat quality
incorporate the needs of orangutans through

sensitive protection
agriculture and
development
orangutan
measures
the
for
enforcement
of
palm
oil
Section
38
including
of
governmental policy
land
plantations
regions,
habitat
Conservation policy should be integrated into
for small-scale
control of
strict
de-
in
graded areas through enrichment planting.
Rehabilitation and translocation.
in
In
allocating
scarce resources, priority should be given
the
conserving
the Wildlife
populations
to
situ,
as
care of former captives ex
situ.
wild
in
Conservation Enactment.
opposed
Tourism industry. Policies should be adopted
Rehabilitation centers should be licensed and
for
enhancement and development
the
monitored,
of
sustainable and responsible orangutan tour-
ism
in
Sabah, both
to
minimize
its
impact on
conservation
lex
most
Current
situ].
enhanced
be
complement
to
conservation measures
Research.
Current
orangutans should continue
Sabahs
be promoted
and enhanced, especially through
records
made
activities
Orangutan High
site-specific
Public awareness. Awareness of orangutan
orangutan
needs and the legal framework
reserves
for
their
must be heightened, especially

and plantation
in
Viability
January 2004,
require special attention and these
and departments.
policy makers, forestry
conser-
Assessit
was
recognized that several orangutan populations
as
protection
improved and
be
freely available to the
ment Workshop held
carried out at local universities, institutions,
among
methods should be used.
keeping should
During a Population and Habitat
on
to
effective
Record
on-site
[in situ].
research
regulations
pertinent
vation community.
conservation activities ex situ should continue

to
all
should be appropriately assessed, and the
orangutan populations.
Off-site
and
should be followed. Rehabilitation methods
the environment and to enhance the conservation of
to the
were designated
Priority Areas'.
Furthermore,
measures were agreed

populations
(e.g.
in
to
commercial
protect
forest
Sabah Foundation, North Kinaba-
tangan, and Trus Madil, where the priorities are:

Ian
Redmond
A palm oil
managers, and workers.
plantation in
Malaysia.
In
the context of these overall goals, priority actions
would include those
listed below.
Research. There
is
need
for in-depth field
studies to
investigate
of logging
and associated human
further the
impacts
activities
Isuch as illegal killing! on orangutan ecology
and survival
in
unprotected forests;
to
assess
the true role of these habitats for ape con-
and
servation;
to
design forest-management
strategies that could

survival
of
allow the long-term
orangutans outside protected
429
World Atlas
keep under natural forest those areas
to
which the
largest
orangutan
For orangutan populations
in
Lower Kinabatangan
populations
occur;
developed
Deramakot Forest Reserve
in
in
the
in
monitor
commercial forests on
orangutan
to
through regular aerial surveys
trends
to
plans with
ground and aerial surveys;
special attention to orangutans;
enhance awareness
to
ted with the workers, contractors,
and
activities
those
In
develop orangutan-based ecotourism that
to
managers,
will provide
economic opportunities
to
local
communities.
relevant stakeholders.
all
research
protected areas;
campaigns conduc-
vation through education
promote
to
orangutan conser-
of
deal with problem orangutans;
to
monitor orangutan populations through
to
stakeholders, with
relevant
all
ways
connect currently Isolated protected areas
by creating forest corridors between them;
of their nests;
develop and implement forest-management
to
reduce conflicts with agriculture by identi-
fying
population
illegal logging
or any other threats to the habitat;
of
orangutan ecology and survival;

to
enhance protection against
to
on the long-term Impacts
forest exploitation
(e.g.
Tabin
priorities are:
Upper Kinabatangan;
to initiate studies
Sanctuary.
Reserve. Kulamba Wildlife Reserve), the
Wildlife
use reduced-innpact logging systems, as
to
protected areas
in
Wildlife
FURTHER READING
Ancrenaz. M., Gimenez.
I.
0..
Ambu.
L.,
Ancrenaz.
K..
Andau.
new estimates
120051 Aerial surveys give
Goossens.
P.,
orang-utans
for
http;//dx.dol.org/10.1371/journal.pbio. 0030003. Accessed
MacKinnon,
11:
J.R. 119711
The orang-utan
in
Sabah
today,
B.,
Payne,
J.,
Tuuga,
A.,
Lackman-Ancrenaz,
Sabah, Malaysia. PloS Biology 3
in
December
111:
e3.
8 2004.
study of a wild population
in
the Ulu
Segama
Reserve. Oryx
Ul-191.
McMorrow,
Talip,
J.,
M.A. 120011 Decline of forest area
land capability Global Environmental

Rijksen, H.D., Meijaard, E. (19991
Century. Kluwer
Change
Our Vanishing
in
11:
Sabah. Malaysia: relationship
to state policies, land
code and
217-230,
Relative: The Status of Wild Orang-utans at the Close of the Twentieth
Academic Publishers. Dordrecht.
MAP DATA SOURCES

Map
17,2
1.
Orang-utan Status
(20041
in
Sabah: Distribution and Population
Size.
Kinabatangan Orang-utan Conservation Project. Sandakan. Malaysia.

Meijaard.
E.,
Dennis,
Meijaard
Borneo Orangutan PHVA Habitat
R.,
Singleton,
I.
& Dennis
120031 and
amended
(20041
by delegates at the Orangutan
Units:
Composite dataset developed by
PHVA Workshop.
Jakarta. January 15-18
2004.
Singleton,
R.,
I.,
Wich,
Byers,
S.,
0.,
Husson,
S.,
Stephens,
S.,
Utami Atmoko,
S.,
Leighton, M., Rosen, N., Traylor-Hotzer,
eds (20041 Orangutan Population and Habitat
Viability
K..
Lacy,
Conservation Breeding Specialist Group. Apple Valley. Minnesota.

For protected area and other data, see 'Using the maps',
ACKNOWLEDGMENTS
Many thanks
to
Marc Ancrenaz iHutanj. Melvin Gumal
(Wildlife
Parks Board. Singaporel. and Lee Shan Khee (UNEP-WCMC)
AUTHORS
Kim McConkey. UNEP World Conservation Monitoring Centre
Julian Caldecott.
UNEP World
Edmund McManus. UNEP World Conservation
430
Conservation Societyl. Geoffrey Davison (National
for their valuable
Monitoring Centre
comments on
World Atlas
Afterword
apes,
Great
among
are
the
interesting creatures on our planet.
First
of
relatives, w/ith
mere
many ways,
in
most
aLl,
they are
chimpanzees
our closest
differing
1.2A percent of their genetic
makeup. Given
apes provide a window
great
into
our evo-
our
in
to
what we ourselves
than
generally realized. The public usually thinks
is
three
of
great
orangutans], or
apes
maybe
(chimpanzees,
four (the bonobol
that there are at least six species
how our minds work, how we

learn, and how we live together in societies. What
is more, great apes may provide clues for cures
to many human illnesses, especially given that they
often share the same diseases with humans
without necessarily showing the same symptoms.
and no less than 13
In
of
addition, great
apes play a key ecological
the tropical forest systems
as major seed dispersers
in
in
which they
role in
live,
in
three genera,
These include two spe-
taxa.
chimpanzees, two species and
at least four different
delve
further into remote parts of the forest and learn
more about these animals through genetic studies,

is likely that this number will increase even more.
it
Unfortunately, as
we
we
more about them,
learn
are also seeing their rapid
decline almost
everywhere that they occur According
with their nest-building and feeding activities.
Mammal
among
we
kinds of orangutans, and the bonobo. As
often even modifying the architecture of the forest
Great apes are also without a doubt
they are
cies and four to five taxa of gorillas, four taxa of
acting
many ecosystems and
gorillas,
if
paying attention. But recent studies have indicated
lutionary history, as well as breal<throughs into the
understanding
did at an
history.
Great apes are also a more diverse group
living
from us by a
this close relationship, studies of wild populations

of
must be very similar

early stage
to the Global
Assessment, an international
assess the conservation status
of
effort
to
mammal
all
known animals
in
species carried out under the auspices of the
the world. They have played important roles
in
Species Survival Commission
the most charismatic and best
many
different
countries
parts of the world, both
which they
in
live
in
the
and many others that
of
lUCN-The World
Conservation Union, at least three great ape taxa

are Critically Endangered and close to extinction.
from
the
have learned about them through books and films
These are the Cross
- both fanciful and scientific - and, increasingly
Nigeria-Cameroon border, the eastern lowland
through tourist
visits to
which they
still
occur.
important
to
cultures that
the often remote areas
the African
live
and Southeast Asian
with them, with
and legends that compare them
encourage
in
They have been particularly
to
many stories
humans and
their protection by placing taboos on
killing or eating
them, or sometimes regard them
with great fear and awe. Their resemblance to
humans
is
striking to all
who observe them. They
use tools, they show empathy, and some,
chimpanzees, even hunt
in
groups,
in
like
ways
gorilla
from the eastern Democratic Republic
gorilla
of the
Congo, and the Sumatran orangutan from northern
Sumatra:
all
three recently featured
in
list
of
the
Top 25 Most Endangered Primates on Earth. The
Red
List
assessments
and subspecies are
for all the great
still
underway, but
ape species
it
is
virtually
certain that all will continue to be classified as
Endangered or
Critically
Endangered
for
the
foreseeable future.
the
that
River
In
in
other words, our closest
big trouble.
As
is
living relatives
so clearly indicated
are
in this atlas.
i3^
World Atlas
their habitats have
modified
in
many
been destroyed or dramatically
of the
places
in
and several species hang on

fragments. Those that
of
the
in
tiniest
of
have a reasonable
still
of habitat available -
amount
which they occur,
as
the
in
Congo
logging and the associated
bushmeat trade
Inot
subsistence hunting but a commercial trade to

serve a luxury market). What
consumption
apes,
clearly
is
it
of this kind
is
is
more, bushmeat
not just bad for great
human
an enormous
health
hazard as well, with strong linkages having been
between consumption
established
meat and outbreaks
human
of the
of
great ape
deadly Ebola virus
in
in
Well,
some
this atlas. First
need more protected areas
of the solutions
many
local
Africa
of
to
be easy
solutions will not
long-lasting
we who have been working

primates for so many years are optimistic
solutions can be found and that we should be
Nonetheless,
to
maintain viable populations of
indeed
nonhuman
all
all
primates,
people interested
in
bode well
that
able
their natural
number
great apes and the in-
them
for the future.
of
presenting
such an attractive and useable
in
it
by bringing together such an

data on the great apes and
makes an enormous
format,
servation efforts on behalf of the great apes.
living in
with
great apes, and
in
habitats. Indeed, the rapid increase in the

of
the
be done, and
to
This atlas,
protected areas and the great apes
them provide
who share
the
like to offer
my
contribution to con-
congratulations to those
benefits
worked so hard over the past few years
broader
to fruition.
would
who have
to bring
it
in
have demonstrated that protected
We
need
these populations, and
do everything possible
highly destructive
132
many challenges on
needs
enormous body
to
keep close
monitor their
health and viability on an ongoing basis. And
need
finding
that
remaining populations as
areas alone are not enough.

track
much
also need to demonstrate
populations
of
more
environment. That said, recent Ebola outbreaks
West
To be sure, there are

horizon and
we
and other creatures

to
Ebola through consumption
of the
possible. To do this,
that these
like
great ape meat.
and foremost, we
for great apes,
parks and reserves that are well managed and

protect as
diseases
the transfer
creasingly strong organizations working on
populations.
What can we do?

are outlined
of
human
those
for
much from
populations that suffer so
forest
Central Africa - are under heavy pressure from
apes themselves but
great
bushmeat
to put
an end
we
to the
trade, not only for the
Chair,
lUCN/SSC Primate
Specialist
Group
President, Conservation International
ANNEX: GRASP and Partners
Annex
Great Apes Survival Project
GRASP and
Partners
UNEP
Great Apes Survival Project IGRASPI
The
innovative and ambitious project of
is
an
United
ttie
communication platform
Nations Environnnent Programnne lUNEPI and

Educational,
Nations
United
the
lift
and
(UNESCO! with an immediate
Cultural Organization
challenge - to
Scientific
the threat of imminent extinction
faced by gorillas, chimpanzees, bonobos, and orang-
GRASP'S mission
utans.
halt
"to
is
have something
to contribute
those
all
response
individ-
current crisis,
to the
Klaus Toepfer, the Executive Director
GRASP
launched
apes and their
new approach
- a
habitat.
visits, field projects,
Through high-level technical
and National Great Ape Survival
and Southeast Asian great ape range

political lobbying
GRASP
countries,
value
it
adds
has
bringing
states, as well
II
in
donor
a strong case for the
ape conservation
World
Development Type
alliance
made
African
in
and awareness raising
to great
GRASP,
UNEP,
of
save the great
to
Plan INGASPl policy-making workshops
as
national alliance
GRASP
Partnership,
UN
is
truly inter-
a diversity of stakeholders.
Patrons have provided their world-renowned
range states have catalyzed government action

to the crisis.
NGASP workshops
to
have helped
ape range countries develop conservation
great
strategies.
GRASP
funding of
NGO
partner projects
has involved local communities and brought about
immediate successes
in
the
field.
conferences, meetings with key
other forms
solidated the
relevant
of
policy
GRASP
biodiversity
Intergovernmental
GRASP
Partners, and
implementation have con-
Partnership and linked
mechanisms and
it
to
multilateral
agreements. Information and awareness
activities
through such media as television and newspaper

articles,
events
efforts.
Summit on Sustainable
together
unique position as a
among
the plight of the great apes. Technical missions to the
to
uals and organizations, the great apes are on the very

of extinction. In
GRASP activities have helped

GRASP might adopt to address
inception,
this crisis, given its
respond
many
its
who
do so."
edge
Since
define the strategies
expertise and reputations to bring further attention to
have the opportunity
Despite the dedicated efforts of
order to bring the
in
the decline
great ape populations by ensuring that
in
decline of great ape populations to a halt.
publications,
have
raised
documentary
films,
the
of
Partnership at the global
profile
and side
the
GRASP
level.
dynamic
agencies,
govern-
GRASP
aims:
ments, nongovernmental organizations (NGOsI, foundations,
and private-sector
links with
promote
message
its
As such,
makers,
it
to
action, to
interests.
By using close
governments through the UN, GRASP can
is
at the highest political levels.
uniquely placed
to
inform
policy
mobilize and pool resources for effective
ensure
maximum
efficiency,
and
to provide
to
lift
the threat of immediate extinction;
to raise
to
funds for great ape conservation;
develop a global strategy to coordinate
efforts to halt the decline of great
tions
ape popula-
and ensure the long-term survival
of their
natural habitat;
433
World Atlas
to
educate local people and encourage sustain-
community
able
to provide alternative
new and
raising
additional
resources - from
country donors, from foundations and the private
activities;
income opportunities
and
sector,
to
from
mechanisms and
existing
hunting, logging, and mining, such as sustain-
national grant schemes. To ensure the long-term
able agriculture, ecotourism,
conservation of viable populations of the wild great
etc.;
improve the infrastructure of protected
to
apes and their

munity
areas;
improve the capacity
to
government
of
made
the
habitat,
international
com-
the widest sense has to provide effective
and coherent support
wildlife
agencies;
to
in
to assist the efforts
being
by the great ape range states.
exemplify the added value of a UN-facilitated
global partnership, with range state govern-
Contact
ments and NGOs assuming Increasing
Great Apes Survival Project Secretariat
control
over the process.
Programme
PC Box 30552
The endangered great apes share
many
with
millions of people
East Africa and
these people
to
in
live
their habitat
below the poverty
central objective of the
Nairobi
West, Central, and
Southeast Asia. The majority
the welfare of
link
in
line.
The need
humans and wildlife

GRASP Partnership.
Further progress will depend
Kenya
is
Fax: 1254 201
http://www.unep.org/grasp
critically
African Wildlife Foundation (AWF), founded
FOUNDATION
AWFs
continent.
wildlife
in
on
http://www.unesco.org/mab/grasp
and wild lands
of Africa
Ape
for Africa's people.
of Africans in conservation.
of the
Congo
Programme described
for
AWF
has worked
27 years, and
Since
to protect
both the
to protect
inception,
its
mountain
a co-founder
is
AWF
has
means
to
gorillas
in
and funder
of
the
below, http://www.awrf.org
Alliance lApAll provides a forum for the discussion of issues relating to apes. An international coalition of
organizations and individuals working for the conservation and welfare of apes,
to tackle
a*
a conservation organization focused solely on the African
protected endangered species and land, promoted small business growth for African communities as a
International Gorilla Conservation
''on for
is
and ensure a more sustainable future
Rwanda, Uganda, and the Democratic Republic
1961,
Partners
programs and consen/ation strategies are based on sound science and designed
improve livelihoods, and trained hundreds
623926
E-mail: grasp@unep.org
GRASP
AFRICAN
WILDLIFE
20)624163/621234
Tel: 1254
of
the problems they face, both
in
the wild and
in
captivity,
together about 70 organizations and hundreds of Individuals,
organizations
and includes an interactive noticeboard
all
working
for
undertakes collaborative action
it
through specialist working groups.

for apes.
The website
anyone wishing
to
links to all
do something
brings
It
member
for
apes.
http://www.4apes.com
Australian Orangutan Project lAOP) works
species
austrdllan
orangutan
project
in
their natural habitat
preserve orangutan populations
funds
to assist in situ
go straight
636
in
orangutan projects
AOP
to the active
ensure the survival
conservation organizations, as well as

salaries are paid to
to
and promote the welfare
its
in
of all
and the
their conservation
own
projects.
AOP
is
of
both Sumatran and Bornean orangutan
orangutans.
AOP
raises
awareness
intrinsic value of individual
of the
need
to
orangutans, and raises
and welfare work. AOP supports many orangutan

a nonprofit organization staffed by volunteers.
No
volunteers and most services are donated. Therefore a very high percentage of donations
welfare of orangutans and on habitat protection, http://www.orangutan.org.au
Balikpapan Orangutan Society-USA IBOS-USAl

Malaysia and Indonesia on the islands
plight
of
is
dedicated to the conservation of orangutans and their habitats
in
Sumatra and Borneo. BOS-USA also raises awareness on the orangutans'
and funds conservation pro|ects http://www.orangutan.com/
BerggorilLa
& Regenwald
DirekthlLfe (BRDI focuses on the eastern gorillas by supporting projects contributing to their
conservation, for example by providing necessary equipment to rangers and park managers.
BRD
projects for the conservation of certain populations of western gorillas that are particularly at nsi<.
supports public awareness
Bonobo Conservation
addition,
BRD
population censuses, and ecological studies, http://www.berggorilla.de/
activities,
Initiative IBCil
also supports
In
has a mission to promote conservation
bonobo and
of the
its
tropical forest habitat
the Democratic Republic of the Congo. BCI uses a multi-sectoral approach emphasizing stakeholder involvement and
in
BONOBO
addressing the needs of local populations, as well as building capacity of Congolese institutions and NGOs. BCI has
established over a million acres under accords for community-based reserves,
and
for conservation,
is
developing sustainable development projects
ecological surveys and information exchange
leadership
in
awareness about
this
Born Free Foundation IBFFI campaigns

against the keeping of animals
in
of the
in
is
bonobo
It
supports the rangers
Team
habitat, motivating local involvement
who
its
and conservation
habitat http://www.bonobo.org/
of
animals
the habituated eastern lowland gorillas.
It
has been working
It
in
the
in
UK and
Cameroon with
around the Dja Biosphere Reserve, and with MINEF and the Cameroon
bushmeat trade
at sanctuaries at the
education programs
overseas
closely with
to
Crisis
m.
the Democratic
of the
GRASP
partnership with governmental
in
Wildlife Aid
in
Fund
apes
relation to
to
living in
and
care for the orphans of the
Mvog-betsi Zoo and the Mefou National Park, and to provide conservation
mmaa
schools and visitors to the sanctuaries http://www.bristolzoo.org.uk
Makerere University Faculty
itself
in
Budongo
and the surrounding population.
and Nature Conservation, the Royal Zoological Society
of Forestry
Task Force (BCTF)
is
of
effective
^03tf^
a consortium of organizations and individuals dedicated to the conservation of
threatened by unsustainable exploitation for meat,
and implement
in
Andrews University http://www.budongo.org
Scotland, and St
and appropriate solutions
in
to this
Africa
and around the world
'bushmeat
support education and training, engagement with key decision makers
by
crisis',
in
We
managing
government and
help
members
identify
BUSHMEIir ClliaS TASK FORCE
scientific information to
private industry,
and
raising
awareness http://vvww.bushmeat.org
animals
in
need throughout the
a rescue center
in
Central Kalimantan,
Care tor the Wild International (CFTWI) promotes the conservation and welfare
world.
and
orphaned
the Ministry of the Environment and Forests
Forest Reserve. Uganda, as well as undertaking background studies of the forest
public
for
Project IBFP) undertakes research on and conservation of chimpanzees and other wildlife
Budongo Forest
Bushmeat
has provided the core
(MINEFI and the Living Earth Foundation on community engagement and support
wildlife
in
patrol the Kahuzi-Biega National Park
Zoo Gardens IBZGI supports conservation projects
works
and
since 2001. http://www.bornfree.org.uk
and nongovernmental organizations.
It
conducting bonobo and
is
It
also dedicated to supporting bonobo research and
important species of great ape and
for the protection
convert logging concessions
to
habitat.
zoos and circuses and as exotic pets. BFF works with sanctuaries
Congo and monitors
Technical Support
Bristol
critical
working
the Democratic Republic of the Congo. Cameroon, and Uganda, and helped to establish the Pan
African Sanctuaries Alliance.
Republic
in
conservation, and promoting bonobo awareness. BCI
raising international
great apes
programs
is
bonobo
in
It
runs an adoption program, which supports orphaned orangutans
in
of wild
Indonesia, and funds habitat protection and antipoaching initiatives to safeguard orangutans
in
the wild.
http://www.careforthewild.com
biodiversity,
Its
regional
and
to
ICI)
has as
its
programs and funding support
species found
in
mission
to
living
natural heritage and global
harmoniously with nature.
CI's
work, through
ape habitat http://www.conservation.org
Dian Fossey Gorilla Fund Europe IDFGFE) works
to
to live
partners, contributes to the protection of great apes and other
genuinely benefit from their natural heritage. Based
designed
conserve the Earth's
to
demonstrate that human societies are able
integrate
traditional
conservation
to
in
save gorillas from extinction and ensure local people
the UK, the fund currently
manages more than 20
projects
"Dian Fossey
Gorilla
Fund
and research with economic development and education.
http://www.dianfossey.org
435
World Atlas
Dian Fossey Gorilla Fund International (DFGFII carries out extensive gorilla protection and science programs
Rwanda
Center
and
IKarisolê Researcli Center]
for
Conservation Biologyl.
it
the lead partner
is
38 700 km' landscape
in a
in
Congo IKabara Research Center, Tayna
ihe Democratic Republic of the
eastern Congo, protecting eastern
in
lowland gorillas by supporting community-based resen/es and national paries. The DFGFI supports 400
field staff,
antipoaching patrols, and education, community, health, and economic development programs.
http://wvvw.gorillafund.org
Discovery Initiatives
promotes ecotourism,
IDII
work
ntention of enhancing the
discoveryinitiatives
monies from nature

ape habitat
travel.
It
of
vi/orl<ing in
those involved
cooperation with the conservation community with the
arranges quality escorted small group or tailor-made tourist travel itineraries
Rwanda, Uganda. Congo, Gabon, Cameroon, Central African Republic,
in
demand and
local conservation projects by linking into the
in
to great
(Borneo], and
Ivlalyasia
Indonesia http://vvww.discoveryinitiatives.com
'/
Earthwateh
INSTITUTE
Earthwatch Institute engages people worldwide
scientific
in
research and education
field
understanding and action necessary for a sustainable environment.
It
promote the
to
has supported several years
of fieldwork
on
great apes http://www.earthwatch.org
AWF
WWF,
and
FFI
IFFII acts to
working towards the conservation
Filmmakers
for
conserve threatened species and ecosystems worldwide. Together with
of
mountain
is
It
aims
to
to make more,
new audiences to
educate, motivate, and inspire
betteractively
and support conservation, http://wvvw.filmmakersforconservation.org
participate
the GREAT APE
Isee belowl, which
http://www.fauna-flora.org
and helps conservation organizations and filmmakers worldwide
informed, and effective consen/ation films.
SPECIES PROJECT
gorillas,
Conservation IFFCl promotes global conservation through the making, broadcasting, and
distribution of films,
WORLD HERITAGE
Programme
responsible for the International Gorilla Conservation
is
Great Ape World Heritage Species Project (GAWHSP) works
secure the passage of an international declaration
to
and convention designating the four great apes as world heritage species.
the outstanding universal value of each of the great apes, and
It
has been established
response
in
in
recognition of
the unprecedented and imminent
to
threats to their survival http://www.4greatapes.com
Institute for Tropical Forest Conservation IITFCI
research and training
the Albertme
in
Rift
is
the lead organization undertaking conservation-oriented
montane
forests
southwest Uganda,
of
in
particular Bwindi
Impenetrable National Park, Mgahinga Gorilla National Park, and Echuya Forest Reserve. The institute was
established
1991
in
as
semi-aulonomous branch
of
University of
Ivlbarara
Science and Technology.
http://www.must.ac.ug/faculties/tropical_forest.htm
mvjw.
International Fund for Animal Welfare IIFAWl works to improve the welfare of wild and domestic animals throughout
the world, prevent cruelty to animals, and
of both
bushmeat
crisis,
policies that
their regional
afromontane
Programme
forest habitat in
(IGCPl acts
in
advance thewellbeing
practical solutions to the
to
ensure the conservation of mountain gorillas
Rwanda, Uganda, and the Democratic Republic
with activities including research and Iransboundary collaboration.
working
to find
http://wvvw.ifaw.org
and
promote animal welfare and conservation
animals and people. IFAW and other partner organizations are working
partnership with
the
protected
area
IGCP
authorities and
is
a coalition of
AWF,
other local stakeholders
Congo,
of the
FFI,
in
and
the
WWF,
region.
http://www.mountaingorillas.org
International Ranger Federation IIRF). The IRF
ranger associations dedicated
is
an international network
to raising the professional
standards
of
those working at the 'grass roots' of conservation and protected areas.

of great
action to
and community
Members
include rangers from a
number
ape range states, http://www.int-ranger.net
Jane Goodall
<f;fK^%
of national, state,
rangers worldwide and providing a voice for
Institute IJGII has as
make
its
mission
to inspire
and empower people
to take
informed, compassionate
the world a better place for people, animals, and the environment. JGI does this through world class
primate care and research, community centered conservation; and environmental and humanitarian education.
http://www.janegoodall.org
436
NGO
Kinabatangan Orangutan Conservation Project IKOCPl was set up by the Frencti

collaboration with the Sabah Wildlife DepartmentWildlife
of
Sanctuary
It
located
is
Sabah. Malaysia IBorneol The primary goal
in
Hutan
of this project is to
achieve the long-term
Sabah. The project now comprises a highly motivated team of 35 trained
in
1998
in
in
the village of Sul<au, close to the Kinabatangan
staff
viability
from the
IMS
Kinabatangan community http://wwAV.boh.com.my/pl/pubdocM3191
Living Earth Foundation
an international nongovernmental organization that encourages people
is
and work
to learn
together
environmental issues which concern them.
to resolve the
development
great apes and their habitat through
its
Africa
of
actively
involved
is
in
the conservation of
EARTH
ideas into action
ram-forest habitat while conducting long-term research
OF
flora within their habitat.
Indonesia and Malaysia and to raise funds and awareness
in
the
in
objectives are to support
UK and
worldwide.
Indonesia.
In
supports the protection of Tanjung Puting National Park, and other protected and nonprotected areas of
orangutan habitat
critical
its
orangutans and other rain-forest fauna and
conservation work
It
LIVING
Programmes, http://www.livingearth.org.uk
Orangutan Foundation lOF) conserves the orangutan and
OF
carries out environmental education and capacity
partnership with corporations, communities, and governments
in
on the ecology
It
It
also operates a rehabilitation
program
that returns
orangutans
the wild.
to a life in
êwW"
http://www.orangutan.org.uk
Pan African Sanctuary Alliance IPASAI
is
an alliance
of 18
primate sanctuaries from
bring the sanctuaries together for long-term planning, and to
over Africa, designed to
all
9^^
improve collaboration between sanctuaries and
primate experts. PASA also organizes workshops on topics such as the veterinary care
of
orphaned primates.
http://www.panafricanprimates.org
PanEco Foundation
Conservation
the
Sustainable Development and Intercultural Exchange
for
Programme IPanEco-SOCP) works on
all
aspects of the conservation
PanEco Foundation and the Indonesian government Department
of
&
the Sumatran Orangutan
Sumatran orangutans.
[PHKAI formally established the Sumatran Orangutan Conservation Programme ISOCPl.
SOCP
orangutans, their reintroduction
to
form
1999.
also involves the
Frankfurt Zoological Society and the Indonesian Foundation for a Sustainable Ecosystem (YELl. Based
activities include the confiscation of illegal captive
In
and Nature Consen/ation
of Forest Protection
new
in
population
Sumatra,
in
the wild,
research into orangutan behavior and ecology, surveys and monitoring, public education, and habitat protection.
PanEco
itself is
also engaged
environmental awareness
in
a diverse array of other projects focusing on sustainable development and raising
both Switzerland and Indonesia, http://www.paneco.ch; http://www.sumatranorangutan.org
in
Tayna Centre for Conservation Biology ITCCBl aims

It
in
to protect gorillas
and chimpanzees
in
the umbrella of the Union of Associations for Gorilla Conservation and
was formed under
the Tayna Gorilla Reserve
Community Development
Congo lUGADECI. and receives support from DFGFI.
East Democratic Republic of
http://wvirw.gorillafund.org/002_site_ind_frmset.html
Tusk works
to
support wildlife and habitat conservation and promotes sustainable rural community development
across Africa, Tusk supports the Chimfunshi Chimpanzee Sanctuary
chimpanzee release program

development projects
as a key
to
in
and around Virunga National Park and Walikale
in
in
Zambia, as well as an orphaned
Conkouati National Park. Congo. Tusk also funds different conservation and
in
Tus
Congo
conserving both mountain and eastern lowland gorillas and their habitat, http://www.tusk.org
UNEP World Conservation
("lonitoring
Centre
lUNEP-WCMCl
provides objective, scientifically rigorous products and
sen/ices that include ecosystem assessments, support for implementation of environmental agreements, regional and
global biodiversity information, research on threats and impacts, and development of future scenarios for the living
world.
UNEP-WCMC
has produced
Volcanoes Safaris runs
gorilla safaris
in
Rwanda.
the development of tourism,
Park and Nyungwe
in
this atlas in
It
enhance
in
GRASP
the gorilla habitats of
works closely with
local
private sector capacity,
UNEP WCMC
http://vTOrw.unep-wcmc.org
Mgahinga and Bwindi National Parks
in
Uganda and
communities and conservation organizations
votc.^xots
to help
and help communities around Volcanoes National
Rwanda http://wv/.volcanoessafaris.com
Wild Chimpanzee Foundation IWCFl works

habitat throughout their range
while also starting
support of
some
in
Africa.
activities in
to
preserve the remaining wild chimpanzee populations and their natural
WCF
is initially
concentrating
its
efforts in different
West African countries
W:
I
Central Africa, http://wnww.wildchimps.org
637
World Atlas
'-
(WCSl works
WILDLIFE
CONSERVATION
conservation, education, and the
SOCIETY
save wildlife and wild lands through careful science, international
to
management
system
of the world's largest
of
urban
wildlife parks.
,...
working
to protect all four
subspecies
of gorilla
as well as working
to protect
WCS
has been
chimpanzees and orangutans
in
their
native habitats http://wvwv.wcs.org
WWF-The
Global Conservation Organization acts to conserve the natural environment and ecological processes
worldwide.
WWF
involved
is
the conservation of the great apes
in
Programme and through
Gorilla Conservation
its
African Great
in
part through
support
its
to the International
Apes Programme, http://wvvw.panda.org
Zoological Society of London IZSL) aims to achieve and promote the worldwide conservation of animals and their
habitats.
ZSLs Bushmeat and Forests Conservation Programme
bushmeat research
LIVING CONSERVAT
West
in
Africa,
and national park development
is
in
centered on equatorial Africa, and focuses on
Gabon and the Democratic Republic
of the
Congo.
http://wftww.zsL.org
ENVIRONMENTAL CONVENTIONS
Convention on Biological Diversity ICBD) aims
and equitable sharing
P
CBD
at conservation
from the use
of benefits arising
and sustainable use
of genetic resources.
areas, which seeks to establish effective networks of protected areas, and

biological diversity,
which promotes sustainable harvesting
enforcement, are of particular relevance

the
to
of
The
its
of biological diversity,
CBDs program
of
expanded program
and the
fair
work on protected
of
work on
forest
timber and non-timber forest resources and forest law
the conservation of great apes. All individual ape range states are Parties to
CBD. http://www.biodiv.org

species
of wild
All great
apes are
listed in
in
Endangered Species
becomes or remains
fauna or flora
Appendix
of CITES.
of
Wild Fauna and Flora ICITESI aims
to
ensure that no
subject to unsustainable exploitation because of international trade-
Trade
in
specimens
of
these species
is
permitted only
in
exceptional
circumstances, and for non-commercial purposes, http://www.cites.org
Convention on Migratory Species ICMSI aims

their
range CIvlS
is
interested
in
to
conserve
between Uganda, Rwanda, and the Democratic Republic
World Heritage Convention IWHCI
is
heritage. Countries
to identifying
nominate
of the
common
Congo, http://www.cms.int
if
of
outstanding universal
heritage of humankind. Countries which have ratified this
and safeguarding the world's most outstanding natural and cultural
sites in their territories
are inscribed on the World Heritage List

of great apes,
and avian migratory species throughout
based on the premise that certain places on Earth are
value and should therefore form part of the
agreement are committed
terrestrial, aquatic,
the conservation of the mountain gorilla, which crosses the mountainous border areas
they
which they believe are
meet the
criteria.
A number
of
of
World Heritage
these sites are
quality,
and these
critical for the survival
http://www.unesco.org/
DONOR GOVERNMENTS
These are governments
that offer funding for the conservation of great apes.
GREAT APE RANGE STATES

The range states consist
of 21 states in Africa
and two
in
Southeast Asia:
Angola
Equatorial Guinea
Ivlalaysia
Burundi
Gabon
Nigeria
Cameroon
Ghana
Rwanda
Guinea
Senegal
Congo
Guinea-Bissau
Sierra Leone
Cote
Indonesia
Sudan
Liberia
Uganda
Mali
d'lvoire
Democratic Republic
of the
Congo
PRIVATE SECTOR
GRASP acknowledges
involvement
438
in
the importance of the private sector and seeks to encourage private sector investment and
the conservation of great apes.
Index
INDEX
Note: Page references
bold refer to illustrations and maps;
in
boxes and tables
tfiose in italic refer to
Alas River/Valleyl 85, 200

Albertine
Alchornea spp.
abominable snowman'
Acanthaceae
lyetil
26-7
303
Association Malienne pour
Conservation de
60
in diet
bonobo95, 251
Alismataceae 85
eastern chimpanzee ilO-11
alkaloids 116
western chimpanzee 356, 377,
alluvial forests 175
391, 3 W, 398
108.
4framomivm
Africa
biogeography 37-40
Ammann,
Karl
27
Anacardiaceae
35,
activities
233-5
conflict
named African
298-9
legislation
Conservation of Nature and
action
African Protected Areas Initiative
African Wildlife Foundation lAWF)
A3A
aggression
166, 167
chimpanzees 63-5
gorillas 100
animal welfare movement 243
Annan,
107,208
87,
Anonidium mannii 87
397
in diet 60, (57,68.
weaver
110-11, 134
CAR
word 14
lApAll 434
87, 108, 116.
117. 123,
7% 321,
326
Mbeli 113-14, 123, 124
Balikpapan Orangutan Society-USA
IBOS-USAI 435
133
arboreal
124
Iboundji 114
Maya Nord
10-11
'ape', origins of
Bambama-Lekana
lifestyle
National Park
325
as impetus for evolution 20
bamboo
orangutans 155
Bangassou forests 315, 317

Bantu peoples 93
Artocarpus 35
Arundinaria alpina Yushania alpina,
{
bambool39,
Asia,
39, 133, 134
banteng IBosjavanicus] 171
Arecaceae 107
subsistence 181,413.417
Aka people314-15, 317, 323
Bafing-Faleme Reserve (proposed!

374, 377
gibbons 209
194,213,368
35, 164
Bating Biosphere Reserve
fca/sl07, 113-14, 124, 126
aging, research using apes 26-7
slash-and-burn/swidden 374-5,
61,71
ants
Apocynaceae 59-60,
405-6
24,
Bai Hokdu,
Ape Alliance
and habitat loss 78, U6, 222,
baboons
anthrax 330
eastern 136, 137-8
agriculture
B
Ba'Aka lAkal people 314-15, 317, 323
IproposedUS, 374, 375
Kofi A. 7
western 114
Aglaia spp. 35, 164
284
raising 280-1,
see a(so public education
Baccaurea spp.
295
Annonaceae
lAPAII 250
and conservation
247 295-8
threats to great apes 220,
Law Enforcement
and Governance 273-i
shifting
55,
120,294,295
56, 105, 106,
countries
Bornean orangutan
apes
future conservation strategies
African Convention on the
Natural Resources lACC) 250
434
awareness
distribution of great
see also
Zoological Parks and
Australian Orangutan Project lAOPI
293, 295
250-1
war and
151,
of
Aquaria (ARAZPAI 263
208
background and economy 230,
agreements and
African Forest
Australasian Regional Association
Angola 293-9
great ape distribution 286-7

regional
Austin 46-7
Association lAZAI 263
spp, 85, 107, 108, 109
116, ]32,
133
Attenborough, David 25, 280
American Zoo and Aquarium
;n, 380
et
377
Asteraceae ICompositael
196, 197
Mountain Wildlife Sanctuary
la
Faune
la
de I'Environnement lAMCFEl
altitudinal limits 35, 175, 178, 185,
Adapoidea 19
la
Protection des Oiseaux
16
alchornidine 116
algae,
John 266
Association Burundaise pour
alchornme 116
132, 133
action plans 250-1
Afi
Aspinall,
Highlands 39
Rift
133, 134
see Southeast Asia
baobab tree 69
barbets 171
Barito Pacific
Timber Group 222
^9
World Atlas
conservation 50-1. 93-6, 253
Barito Ulu forests, Central
Kalimantan 175
action plans 95, 251
Baselta
33
43, 53, 83,
Basellaceae 133
Zoo Gardens 310. 312. 435
bronchopneumonia 235-6. 386-7
Budongo Forest
88
distribution 50, 83-5. 333,
334
Project 435
Budongo Forest Reserve
61. 62. 71
evolution 43-4
buffalo 135
Batak people 226, 227
habitat 45, 85
Build Environment for Gorillas
Batang
human
195
bat. fruit 163, 171,
National Park, Malaysia
Ali
Batang Gadis National Park 273.

422
Battell,
Andrew U-15
bear, Himalayan black 26
Kalimantan 214
see traditions and beliefs
benefit stnaring
258-9
Berggorilla
sanctuaries and rehabilitation
Burkina Faso 54,
Borassodendron borneensis 1 64
& Regenwald
Betung Keritnun National Park
and feeding
Ghana
unknown ape
Bia National Park,
79.
355
ranging 164, 166
Bili-Bondo.
27,
334
response
'bimatunsm', orangutans 56-7
social 166-7
vocal 167-8
release of apes from 370
CAR 317
Hi
birds 171, 173,
7(5,
208, 209. 212
CAR
Blumeodendron
and
Bofi people
sp.
207
diet 31
beliefs
and traditions
227
Malaysian, seeSabah; Sarawak
Foundation 182.252,422.423
Bossou
435
bonobo [Pan paniscus]
94. 339,
(BCII
33. 45
captive breeding 263
Congo 121-2,323,324,325
control
measures 246-7, 260-1,
DRC50,
92, 93,
122,335.338
Equatorial Guinea 122, 228-9,
344-5. 346
factors encouraging 50. 237.
268
65
Thomas
16
Gabon 122,349,352
Brachystegia 40
Ghana 355
brain size 23-4
gorillas 101. 102, 121-2, 143-4,
Brazzaville Process 244-5, 351
behavior 45-9. 85-90
and feeding
58, 60,
Bowdich,
44. 78. 368, 374,
398, 402
283,311,325,346
341
317
Bombacaceae 69. 163. 187

Bonobo Conservation Initiative
315, 317
consequences 68
Bososandja Faunal Reserve
33
338
chimpanzees
Born Free Foundation 410, 435
blowpipes, use of 198
50,
Cameroon 122,307,309,311
Borneo Orangutan Survival
see reproductive rate
birth rates,
diet
179-80
populations 181
226,
seed dispersal 171
size,
bonobo
human
BirdLife International 310
body
Angola 295
subspecies 154, 161
biogeography and ecology 33-7
bipedalism 23,
Task Force (BCTFI
bushmeat hunting 226-9
Borneo
359
Crisis
population 180, 787, 218, 219
distribution
Biosphere Reserves 3^9. 352. 355.
344-5, 368
245, 435
evolution 153
biomedical research 27-8, 370, 398
markets 226
scale of trade 226, 228-9
Bushmeat
endangered status 179
37-40
bushmeat
taboos and traditions 227. 338.
289,418,419
Southeast Asia 32, 33
302
historical
Sumatran
orangutan 154, 155
biogeography
Africa
167
distribution 161. 162, 179-80,
and conservation
Burundi 301-2
differences from
of infrastructure
304
bushbuck61. 135
development and reproduction
development 236-7, 239
Bururi Forest Nature Reserve.
biodiversity 37
261,438
55,
threats to great apes 78, 220,
disturbance
to
binturong Ibearcatl 195
Convention ICBDI 216, 249-50,
apes
action 247. 302-4
168-70
Africa 37
299, 301
legislation
ecological role 170-3
181,255
impact
161, 163-4,
170
37,
56-7,300,301-2
behavior and ecology 161-74

diet
288
pygmaeus]
DirekthilfelBRDU35
56,
Burundi 299-304
Bornean orangutan [Pongo
Benin 54, 56
191,271
Bunopithecus, see gibbons, hoolock
threats to 223-4, 227, 337
60.
in diet
Bukit Tiga Puluh National Park 154.
reproduction 88-9. 218
339-40
bearcat (binturong! 195
beliefs,
218
lifespan
populations 50, 83, 90-2, 218
28;
IBEGol 151
Bukit Baka National Park.
physical characteristics 83
BBC Worldwide
bees,
92-3
i28
IM
Bristol
differences from chimpanzee
bark, as food 164, 170. 187
BriUantaisia 133
335,338.381
Brazzaville Zoo 323
Liberia 368
breeding, captive programs 263-6
and logging 256-7. 324
Bridelia sp. 737
orangutans 178. 198
Index
trapping and snaring 78, MS,
22S-9, 317, 323, 338. 344. 386
use
of
firearms 237, 317, i02
cultures 66, 67
CARE
diet 33. 44.
International 388
Busfimeat Project lUSI 310, 311
Care
Bwindi Impenetrable National Park,
carnivory
Uganda
98, 99, 102, 132,244,
261-2,274
conservation activities 149
gorilla populations
and human contact 65
bonobo
hunting 58-9
45, 85
infanticide
chimpanzees
Interactions with other
44, 58-9, 60-1.
animals 71-2
Castanopsis
gentitii
nest building 69. 71
ranging 44-5. 61-2
87
response
165
sp.
censuses, ape populations
Center
of
Research
Sciences
Cabinda province, Angola
56, 105.
in
for
social 59, 62-5. 349. 351
Natural
DRC ICRSNI 339
Central African Regional Program for
Cameroon Highlands 39
Cameroon 39. 305-13
background and economy
56. 105. 106.
legislation
apes
55.
56.57. 106. ?20, 305-7. 308

apes
55.
and conservation
action 79.
and conservation
action 79.125.247.249.
mapping
US. 2473-17-19
lifespan
forest loss 121. 315.
Aid Fund
ICWAFI
83,88
release of captive 74-5, 79, 253,

267. 2(58-9, 271,326. 356. 370
reproduction and development
Forest Initiative 245, 274
48-9, 65, 68, 218
research50-l,57, 58-9,
Netw/ork IRAPACI 351
taxonomy
chimpanzee IPan
,5,5
53. 56
threats to 27-8, 75-9. 398
troglodytes troglodytes]
disease 74. 78-9. 323-4, 330.
distribution 54-5, 56, 295, 306,
311-12
changes 225
populations 49, 73-5, 218. 240
320
Central African World Heritage
central
of
2/8
physical characteristics 43, 53,
threats to great apes
Central Africa Protected Areas
309-12.317
threats to great apes 121. 122.
220 223
Cameroon Wildlife
habitat 39-40. 44. 57-8
120,315,316
hunting 122.315.317
312-13
legislation
distribution 54-5. 56-7
evolution 23-4. 43-4
314-15
305
people 258, 364
conflict with
species action plans 251. 394
background and economy 230.
230.
use 67 68-9, 361
conservation 49-50, 79-80, 253
Central African Republic ICARl
Calamoidae 33
tool
captive breeding 263-6
314-19
see vocal behavior
to habitat
disturbance 71
124. 141
the Environment ICARPEI 274
125.295,298
Caesalpiniaceae 38. 40. 107, 108
calls,
language acquisition 46-8
Sumatran orangutan 187

Carpodinus
tourism 410
64-5
Bornean orangutan 164
Carpenter. Clarence Ray 24
140. 141
58-61
ecological role 70
Wild International 435
for the
71-2
future researcfi needs 41
mountain
communication 68
gibbons 213-14
Carduus nyaanus 33
307
402. 407
Cameroon 315, 316

Congo 321, 322
habitat loss/logging 76-8
CAIvlPFIRE 281
camphorwood 165
Equatorial Guinea 342-4
live
Camp
Gabon
268-9. 355, 359. 398. 402
Campanulaceae
V6
Camper, Peter 15-16
Leakey 168
hunting 78
348, 350
releases/relntroductions 266-7,
270-1
bonobo 339-40
chimpanzees
74-5. 79. 253,
267.2(58-9,271.326,356,
eastern chimpanzee;
threats to 323
Nigeria-Cameroon
chimpanzee; western
Centre International de Recherches
Medicates de FranceviUe
chimpanzee
74,
Chimpanzee
126
370,413,415
Cercicopithecus spp. 60. 61
gibbons 213-14
charcoal burning 405
gorillas 126, 271
'charms', apes as 226
orangutans 180. 264. 267.
cheek pads Cflanges'l
270,271,422
79, 247,
see a(so central chimpanzee;
genetic studies 56
captive animals
animal trade
Rehabilitation Trust
272
Chrysobalanaceae 58
Chrysophyllum [Gambeya]
154, 157-8,
85. 108
CITES, see Convention on

International Trade
188-9, 190-1
In
research 25-8
chest-beating displays 143
Endangered Species
welfare 243
chimpanzee IPan frog/odyfesl

behavior 44-9. 57-73
Fauna and Flora
see also pet trade
access
captive breeding
bonobo
247,
263
chimpanzees 263-6, 271
to
water 57
coexistence with gorillas 41

135. 137
civet,
of
Wild
arboreal 173
climate change, post-glacial 37-8.
154
climbers,
woody 170
641
World Atlas
Clitandra 133
Club des Amis de
la
Nature 311
Clu5iaceae35, 37, 39, 132.208
56, 105, 106,
226, 246-7, 438
chimpanzee
41, 135, 737
Cola spp. 116
and conservation
collaboration, intergovernmental
habitat loss 121,
317,325
hunting 121-2,323
Congo
Basin, regional
agreement
56, 57,
Congo
black-and-white 89
Conkouati-Douli National Park,
90
61, 71,
Thollons red 89
River 83
Congo
407
and bushmeat hunting 237

1
73
chimpanzee 49-50, 79-80, 253
Resources ICAMPFIREI 281
communication
effectiveness of 7?, 80, 123,
284
gorillas 101-3
crop raiding 143, 364
western 123-5
cross-border collaboration, see

transfrontier conservation
Cross River
integrated approaches 259-60,
gorilla 142
Sumatran orangutan
280-2
193
distribution
Bornean 182
ecology 99,
stakeholder groups 277

field
258-60,273-4,281-2,283,
285,313
Comoe
282-3
and chimpanzees
135
taxonomic history and status
116, 132,
133
ICII
273,
translocation 270-1
369-70, 435
Critical
Ecosystem Partnership
Moist Forest Ecosystems
244,310
Africa,
in
Central
see ECOFAC
between great apes 349, 351
189,
190-1
wildlife
see a/so aggression; human
Convention on the Conservation of
143,258
cryptic
79,230-2,380
apes 26-7
cultures
chimpanzees 66-7
consortships
Sumatran orangutan
31,
of
conflict
Cross River Gorilla Project 31

Cross River National Park, Nigeria
Conservation and Rational Use

Forest Ecosystems
Conference on Central African
Congo 320-7
70
threats to 220, 307, 309, 380-1
Fund 370
Compositae lAsteraceael
conflict
382
chimpanzee-human 393
between people and
70-7
population 101, 709, 770, 27S
transfrontier 125, 255, 283, 299,
gorillas
studies 777
305,310-11,317,325,356,
National Park 330
competition
U2
strategic priorities
70,305,307,308,
380
Sumatran 202-3
gorilla 119
gorilla [Gorilla gorilla
diehli] 109-11
conservation 7?/, 312-13, 381-2
orangutans 158-9
189, 192,
249
crocodiles 168

language [human! i6-8, 91
Site
178,427
financing of 274-5, 276, 283,
eastern 148-9, 151
western
threats to great apes 220, 330
Crocker Range National Park 175,
Cross River 312-13
chimpanzees 68
61, 62, 65, 68,
330
World Heritage
bonobo 90-
mountain
72, 79, 328,
ex situ 263-6
Communal Areas Management

Programme for Indigenous
Park
Tai National
258
125,254
109, 133
and conservation
action 79,247,330-1
93-6
conflict with people
Combretum nigrens
54,
331
legislation
50,
apes
328-30
74-5, 325
conservation
bonobo
coltan mining 144, 236-7, 257, 335,
Commelinaceae
328
black 229
redA4, 5g-y,
340
Cote dlvoire 328-31
244, 274, 324, 351
Miss Waldron's red 355
94
corridors, green/forest 125, 258,
220,324
Congo Basin Forest Partnership
Angolan 90
91,
438
250
colobus monkeys
246-7
Convolvulaceae 133
disease 123,323-4
125,255,283,299,305,311,
Project 66
28,
conventions, international 246-50,
action 79, 247, 324-6
Chimpanzee Culture
Appendix
MIKE program
legislation
of
Wild Fauna and Flora ICITES)
120,321-3
326-7
coexistence, gorilla and
Collaborative
55,
coal mining 181
Endangered Species
in
apes
Cluster Mountain World Heritage

Sites 2A8
Convention on InternationalTrade
320-1
ICBDI246, 249-50, 261,438
human, see
traditions
and
beliefs
Sumatran orangutan 792-5

Curcubitaceae 133
Cyclosorus dentatus 85
Cynometra alexandri
Migratory Species ICMSI 249,
Cyperaceae
438
cytochrome-c
69, 85,
71
109
'clock' 19
Index
gorilla
adaptation of bearded pig
tourism 150
human
beliefs
and traditions
227
Dahomey Gap 37
and conservation
legislation
action 79. 93-6. 338-40
Danau-Sentarum National Park
threats to great apes 92-3, 335,
775, 181
Sumatra and Borneo 33-7,

153
mast
fruiting 161, 163, 165. 197
discovery, of great apes 13-17
337-8
Darfur235
163, 165
species richness 34
Darwin, Charles 16-17. 24
deforestation/logging 92, 93,
Discovery Initiatives
Darwin
220 223-4,
disease 235-6
Initiative
369
Dayal< peoples 178, 425
human
deaths
234,
infant
apes
65, 88-9,
U4,
parl< staff
relief
conflict 50, 92, 144-5,
236-7
U5
355
deer
257-5,232,235,236,321,
World Heritage Sites 249
327, 330, 348, 352, 402
Congo
Kingdom
of
parasites 79, 147
238-40
chimpanzee habitat 76-8, 402
Developpement d'Alternatives au
Braconnage en Afrique
92, 93, 220. 223-4, 335,
Centrale [DABACI 260
337
transmission from
264. 407
yaws 123
Dialium
Ghana
diamond mining 396, 397-8
Africa
Dian Fossey Gorilla Fund Europe
questions 287-9
220. 355
gorilla habitat
eastern 145-6
08
see also under each great ape
Dian Fossey Gorilla Fund
Guinea 359
International 149, 151,337,
Guinea-Bissau 220. 364
388, 436
Indonesia 180-2
bonobo
Liberia 220. 368

177.
181-2
45,
33
33,
58-61
DNAanalysesl9,
34-5, 133, 163-4,
gorillas 70. 98-9, 221
Senegal 393
eastern lowland 134
Sudan 402
Cross River
mountain 133-4
IDRC) 37,
materials 31. 59,85,98, 133,

of
the
Congo
144-5,234,236-7,332-41
332-3
apes
55,
288-9, 333-5, 336
research 85-8, 94
340-1
382
Dryobalanops aromatica 165
187,208
Drypetes
sp.
Duboscia
1 1
164,
Bornean
161,
Sumatran
163-4
185, 187,
and seed dispersal
757
duikers 44. 60
196-7
apes 20
70, 71, 81,
112, 170-2, 194
digestion 31, 70,98
Dipterocarpaceae 34, 208

forests 33, 161, 163, 174. 175,
196-7
and Breeding
Centre, Nigeria 267, 272,
orangutans 34-5, 154-5
role in evolution of
56, 105, 120, 130, 131-2,
field
Drill Rehabilitation
leaves and herbaceous
50, 83, 84, 92,
of
the Congo
western 107-11. 116
Democratic Republic
53,56
DRC, see Democratic Republic
1 1
Sumatra 199-201
see also logging
154
43,
dogs, hunting 195
Tanzania 223-5. 413
Dendrocnide spp. 187
43, 53, 56. 97.
chimpanzees and bonobo
orangutans 154
gibbons 207, 208, 209
223-5
79.
gorilla 97
770
Rwanda 388-9
Cameroon
123,307,309
31,
85
chimpanzees
flexibility
orangutan habitat 168-70
satellite data
under country name
and body size
moist forests 158-9, 168-9,
species/subspecies and
Dja Faunal Reserve,
diet
Kalimantan 180-2
apes
286-7
Southeast Asia 287
257, 388, 435
western 120-1
123,
veterinary assistance 147, 149
Equatorial Guinea 345
humans
126, 147, 750,235-6,262-3,
development
Cote divoire 328, 330
prevention measures 147, 352

respiratory 147, 330
see also reproduction and
Congo 324
DRC
323-4, 348, 352, 407
202 236-7,
infrastructural 126,
219-20
gorillasl02, 122-3, 126, 146-7,
measles 147
economic 198-9, 236
and agriculture 328, 405-6

causes
of
IPigafetta) 14
development
deforestation 10, 219-20, 253
74, 78-9, 323-4,
330, 348, 352, 402, 407
unknown ape 27
Description of the
mouse 171, 172

sambar 172
436
Ebola virus 78-9, 102, 123,
338
U1
(Dll
anthrax 330
chimpanzees
hunting 50, 92, 93, 122,335,
119
silverback gorillas 138.
debt
335, 337
black-fronted 135
blue 61. 229
Durban Process' 257

Dur/ospp. 163. 164. 171. 178
Durrell Wildlife Conservation Trust
311,346
Dzanga-Ndoki National Park

122,
107,
125,315,317-18
U,2
World Atlas
Dzanga-Sangha Dense Forest

Special Resen/e
79,
125,315.
Ebo Forest 307. 313

Ebola virus 123. 231-3 235. 236, 402
chimpanzee5 78-9, 323, 330.
317
Dzanga-Sangha
342
348, 352
project 125
102,323,348.352
gorillas
eagle, black 168
Earthwatch
Institute
436
Ecosystems
in
310.319.325,345,351
346-7
legislation
and conservation
action 79. 247. 345-6
threats to great apes 121. 122.
CAR 319
220
268.
344-5
Congo 325
Esso 256, 309
behavior 62, 6A-5
Euphorbiaceae35. 107,
conservation 338-9, 387-8,
Gabon 351
troglodytes schweinfurthii)
402-4
164,
ecological role of great apes 40-1
action plans 410-11
bonobo 87
distribution 54-5, 56-7, 300,
Eurasia 21-2
European Association
as competitors 41
116. 137.
70 208
of
Zoos and
Aquaria (EAZA! 263, 264
301,315,316,322.333,334,
forest structure 40-1
384, 385-6
gorillas 112. 115, 135
Sudan 400, 401
orangutans 170-2, 194
European Parliament 264
Tanzania 413,414
seed dispersal 70.71.87. 112,
European Space Agency 151
Uganda
405, 406
European Commission 201. 231,

274, 370
European Union
170-2. 194
Economic Community
genetic studies 56
African States
threats to 323, 335-8, 386. 387,
328. 366
of
West
lECOWAS]
bonobo 43-4
ecotourism, see tourism
137
education, public 123, 264, 271-2,
133-4
gibbons 207
282-3,285.311,313
nest building 141
Elaeis guineensis
ranging behavior 134
(oil
products 280-
conservation and research
elephants 92. 135, 194
148-51
El
distribution 129-32
10,287
see endangered status
Nino Southern Oscillation (ENSO!
Fagaceae 164
163, 165.220
physical characteristics 97-8
endangered status
Fang people 227. 229
populations 140. 141-3. 151,2/8
research, gaps
chimpanzees 75-6
in
know/ledge
Red List 9]. 217-18.
eastern lowland gorilla IGorilta
behngei grauen)
Sumatran orangutan
2W
333, 336
ecology 98
habitat 132
population 101. 140. 141-3, 151,
199. 217.
240
reproduction 100. 139. 141
foraging behavior
bonobo 48-9, 86-7

coalition formation 47-8. 86.
138-9
Sumatra and Borneo

El Nifio
33.
34
Southern
Oscillation
environmental impact assessment

(EIA!
68-9
diet;
chimpanzees 62-3
Africa 39
ENSO, see
67.
female behavior
endemism
DRC
use
see also
218
134
threats to 143-8
tool
Cross River 101, 110
tourism 149, 150
distribution 130,
feeding
gorillas 101
threats to 143-8, 150
2<5-7.
151,257.310.369-70,436
gibbons 214
103
subspecies 129
218.
risk,
427, 429
vocal behavior 142-3
diet
extinctions
local
plantations 158. 181, 194,417,
social 135-9
153-4
ex situ conservation 263-6
palm! 60, 71.
158
reproduction 139, 141
21, 22-3,
primate origins 19-21
assessment! 245-6
infanticide 136, 138
humans 23-4
orangutans
EIA (environmental impact
ecological role 135
122. 126.274.
chimpanzee 23-4, 43-4
274-5. 276. 283, 284
coexistence with chimpanzee
diet
ECOFAC program
evolution 17-24
financing of conservation
behavior and ecology 132-41
121, 255, 310, 359,
381.389
310.319,325.345.351
economic factors 230-2
400, 402, 405, 407
eastern gorilla [Gorilla benngei]
i>U>
of Forest
Central Africa! 122, 126,274,
eastern chimpanzee IPan
55,
(Conservation and Rational
Use
apes
56.74-5, 106, 120,342-4
research 327
ECOFAC
245-6
Eocene epoch 19
gibbons 211
gorillas 99-100. 117. 136.
138-9
orangutans 158, 166, 188, 189,

190-1
Rcusspp.
35, 58. 137. 155. 163-4.
187.207
Index
field
seed dispersal 171
changes with
strangling 34. 187, 196
community 273
studies 24-5
194,397,427,429
114,
fuelwood 146. 405

Fulani people 376
dipterocarp 34-7. 153. 161. 163.
ecology 393-4
196-7.208
conservation-oriented 272-3
adaptation of bearded pig
Û. 113-U.
gorillas 24-5, 105,
163. 165
Gabon
dry 39-40
318
language acquisition 46-8

medicinal value
of
foods
U6
ecological roles of apes 40-1.
25, 182
70.71. 87. 112. 115. 135.
tool use 67.
68-9
170-2.194
Moraceae
spp.;
losses
legislation
and conservation
action 79.
125.247 351-2
protected areas 79
lowland ram 38
436
apes 102.
105.106.120.348.350
forest fragmentation
Filmmal<ers for Conservation 281.
348
see deforestation;
of.
348-52
land-use planning 420-1
Fifi225
57.
dry/savanna 40. 45. 58. 107
orangutans
see Ficus
Dipterocarpaceae 34
processing 59-60, 67,

189-90. 192
conversion to plantations 181,
57, 58-9, 66. 67.
68-9, 79
figs,
fruiting.
fruits,
Congo 320-1, 324
bonobo 85-6, 91.94

chimpanzees
altitude 35
Fimbristylis spp. 109
Marantaceae 107.321.325
disease 74, 123,348,352
finance, for conservation 274-5,
montane
hunting 73-4, 122
276, 283, 284

fire,
see forest
32. 33. 35. 36. 60. 175.
logging 40, 73-4, 121,220.
301
restoration 257-8
fires
223.348-9.351
secondary/degraded 40. 107.
firearms, use of 237, 317, 402
76?- 70. 175
flanged' males 154, 157-8, 188-9,
galago. Thomas's 72
Galdikas. Birute 25
Southeast Asia 33-7. 174-8
Galium ruwenzoriense
flying foxes Ifruit batsi 163, 171, 195
sustainable use of 257
gallery forest. Burundi 301. 302
Food and Agriculture Organization
swamp/peat-swamp 38-9.
190-1
Nations IFAOI
of tfie United
85.
219,245,386
;7(5-7 318.
availabilityof 31,33, 34-5, 38,
Gambia
321.323
processing 59-60. 67. Hi.
158-9. 168-9. ?77 181-2
Garcinia spp. 35. 178
see also
fossil
genome comparisons 24
German Development Credit
remains 18-24
Agency 231
early primates 19-20
foraging behavior
early simians 20-1
bonobo 45
chimpanzee 44-5. 70
eastern gorillas 139
Front for the Liberation of the
western 111-12
Enclave
orangutans 34-5. 155. 163-4.
185,187,196-7
of
Cabinda [FLECI
chimpanzees
353. 355
44. 59-60, 70
eastern lowland gorilla 134

39, 220,
420
gibbons 207. 208
56. 354.
impact 37, 40
gorillas 98-9. 133, 134
Southeast Asia 37. 159, 181,
orangutans 161. 163.
199-200,220
protected areas 79
fruit
eating,
Governance IFLEGI process
masting 34. 38. 108. 161. 163.
250
165,
Africa
38-40
fruit
see frugivory
bats 163. 171. 195
gibbons
13.
16.22
agile206. 209. 211
197
seed dispersal by apes
forests
and conservation
action 79, 247, 355-6
Law Enforcement and
Forest
54,
356
legislation
and seed dispersal S7 194
forest fragmentation 121, 126,
apes
355
170, 188.
196-7
201,420
191
261. 353-6
frugivory
forest fires
Sumatran orangutan
Ghana
295
113-U
bonobo 88
415
137
forest clearings 107,
gestation period
Frankfurt Zoological Society 389.
gibbons 208. 209
causes
Nigeria49. 380. 381

genital rubbing 48. 87
apes 21-4
diet
gorillas 134.
gardens' 178
Gashaka Gumti National Park.
389
sharing 61
85,
288
gap analyses 282
Fossey. Dian 24-5. 135. 242.
189-90. 192
56.
disturbance and drainage
257
medicinal value 116
133
108
Forest Stewardship Council IFSCI
39
Gambeya [Chrysophytlum]
102. 109. 119. 174, 175,
foods
39,
87,
194
behavior and ecology 171. 173.

174.
194-5.207-11
^5
World Atlas
212-U
conservation
gorilla;
2U
crested206, 210, 212,
Granada International 281
habitat 207-8
GRASP, see Great Apes
2U
hoolock 206, 209,
Gunung Palung National Park
2U
Great Apes Film
281
Initiative
Great Apes Project 311
lar2A, 205, 206, 207, 210
Great Apes Survival Project
(GRASPI 235, 243-4, 275,
Mullers206. 209. 210, 211
290-1,326.340,433
pileated 206
predation by great apes 187
siannang 19^-5. 210, 214
eastern lowland gorilla 132-3
(GAWHSPI 436
grooming
Gilbertiodendron dewevrei 1 08
bonobo 46-7.
86. 88
chimpanzees
63,
Gishwati Forest Reserve 388
254,274,297.319,356,369
globally significant biodiversity area
IGSBAI system 356
western
Primates (HELP! 74-5. 322.
Gombe
assessment using
Parl<.
Tanzania 74.
eastern 135-6, 138 139
caused by human
western 115. 117. 118
chimpanzees
field
studies 58-9.
imaging 223-5, 413
central 324, 345
eastern 402, 405-6
Gombe Stream Research
Centre
gibbons 213, 214
61,
gorillas
71-2
eastern 145-6
mountain 405-6
07
Guinea 357-61
western 118. 120-1,324,345
24-5
Goodall. Jane 24-5. 29-30. 57. 58.
244. 281
236-7, 238-40
gorilla
220
habituation
360-1
bushmeat 226
conservation concerns 101-3
116.221
differences between species
97-8
and traditions
227
ecology 98-9
and conservation
247
255, 359-60
Guinea-Bissau 227 362-5
endangered status 101
chimpanzees
41, 112, 115, 135, 137

362
apes 362-4
forest losses 220,
research 103
action
gorilla;
39, 132
hallucinogenic plants 116
Halopedia azurea
Hanno
09
iPeriplus] 14, 16
Haumania spp. 107, 109

Haul Bandama, Cote d'lvoire
328,
Haul Niger National Park, Guinea
359
Haul Sassandra forest 328
HDI, see
legislation
resilience to 221
see also Cross River
364
364-5
threats 102
gorilla 105, 123
330
populations 218
taxonomy 97
western
Hagenia abyssinica
legislation
action
chimpanzee 76-7
beliefs
threats to great apes 220. 359
distribution 102
interactions with
human
orangutans 158-9, 169-70 240
see a/so deforestation; logging
forest losses
gorillas
diet 70,98-9,
apes
357-9
word 16
see eastern
infrastructure development
357
'gorilla', origin of
Gorilla beringei,
fragmentation 121, 126,
Salonga 89
Guibourtia
335
199-200,220
guereza59,
65. 66. 79
satellite
149.340
125.
conflict
7-8. 57-8, 76-9
western 330
crowned 60
223. 2S;. 415
chimpanzee
GTZ 122.
guenon
satellite data
223-5.413
gorillas 99
Groves. C.P 27. 53
National
loss/damage 219-20. 220-5
habitat
gibbons 211
see also party formation
Gala Forest Reserves 396. 397
gorilla 102, 105, 106
326
gorilla 117
structure and infanticide 138
239
132-3
gibbons 210
GLGBIO
computer model 236-7.
gorilla
western
Global Witness 310

2
mountain
Habitat Ecologique et Liberte des
65
groups
Global Environment Facility (GEFI
gibbons 207-8
remote sensing 151, 223-5, 413
gingers IZingiberaceael 38, 85, 107,
64
45, 85
Partners 434-8
Project
164
bonobo
chimpanzee 44
2U
Gigantopithecus 22, 23
H
habitat
aims 243-4, 433-4

Great Ape World Heritage Species
taxonomy 205-7
166,
168-9. 175, 181
Survival
Kloss2U
Gironniera nervosa
200
37. 185. 193-4,
211,248,252,283,422
Project
Innoloctil 206. 209,
threats to
Sumatra
gorilla
evolution 207
Javan
UA
western
governments, national 277
and conservation
247 364
Human Development
Index
head-hunting 179, 226

health, perceived value of great
threats to great apes 220. 364
eastern gorilla; eastern
Guineo-Congolian formations 37. 38
lowland gorilla; mountain
Park.
apes 323
HELP chimpanzee
326
project 74-5.
Index
hepatitis,
medical research 27-8,
population pressures 213, 221,
370
herbs,
in diet 85, 107, 109,
133
423-4
refugees 144-5, 146, 332, 335,
Hehtiera elata 187

HIV/AIDS, research 28
197-8
Bornean orangutan
164, 166
rib.k\9-l'i^
by great apes
infanticide
bonobo
gibbons 209
chimpanzees
gorillas 98, 99
71-2, 73, 85
mountain
134
98,
45,
chimpanzees 64-5
85
44, 58-9, 60-1,
apes 226-9
body parts used
197
gorillas 99, 117, 136, 138
infants
orangutans 164, 187

of great
1 1
Sumatran orangutan 187-8,
bond with mother
chimpanzees 65
in
health/
orangutans 158, 167, 188,
medicine 323
Hominidae 13
bonobo
erectus 23
50,
190,191
338
kidnapping by apes 65
chimpanzees330, 335, 359,
floresiensis
27
139
Cross River
137, 191
hooting
gorilla 381
eastern gorillas 143-4, 335,
bonobo89, 91
western
338
gorilla
19
western
221 233-7, 262
Angola 295, 298

Burundi 301
50, 78,
398
143-4, 178, 179,
gorilla
238-40, 324
121-2
Gabon 348-9
sport 179
impacts
see also bushmeat; bushmeat
Indonesia 420
Hutan 182,252
Huxley,
Thomas
of
126,236-7
Sumatra 202
hunting
and bushmeat hunting
in live
268-9
DRC 338
prehistoric 156
173,208,209,212
conflict 101,
trade
infrastructural development 126,
orangutans 178, 198
see also vocal behavior

hornbill 163, 171,
mortality rates 65, 88-9, 119,
381,402
homosexual behavior
human
420-3
hunting
61
and conservation
action247, 249, 267, 270,
hunter-gatherer peoples 174,
chimpanzees 44-5,
western
legislation
368
home ranges
Homo
Homo
386, 396
insects, in diet 60, 110, 133, 164,

14, 16,
17,22
187
Congo 321
Hybophrynium braunianum 69
Congolais pour
la
Cote divoire 328
Hydrocharis spp. 107, 109
Conservation de
la
DRC144-5,
Hydrocharitaceae 107
IICCNI 339, 340, 389
234, 236-7, 335,
337
Hytobates 206
and eastern
and
gorillas
144-5
staff
see also gibbons
gorilla conservation 101
impacts on
wildlife 50, 92, 101,
Hypericum revolutum
Institut
39, 132
hyrax, western tree 72
144-5, 146,335
144, 234, 335, 383,
Leone 396-7
Sudan
235, 400, 402
Tropicale IIRETl 352
Conservation IITFCl 436
385
and tourism 149, 262
Human Development
Index
rankings 230
Development Project
Lowland Rainforest
Iban people 178, 179
259-60.280-1,285
impenetrable Forest Conservation
Project 149
inbreeding 148
disease transmission 123, 126,
In
147, 750, 235-6, 262-3, 2<54,
Defense
of
evolutionary divergence 23-4

habituation of apes 76-7. 105,
Indonesia 417-24
infants killed by
apes 73
ecotourism 262
murders
145,388,389
forest fires 159, 181
112, 115, 135, 137
bonobo 89-90
chimpanzees 71-2
198-9,230,417,419
419
123
between great ape species 41
Bornean orangutan 173-4
Animals - Africa 31
407
259-60,280-1,285
interactions
with other animals
Imperial College London 346
behavior 65
discovery of great apes 13-17
Aceh
development projects IICDPsl
development projects!
contact and chimpanzee
for
integrated conservation and
Iboundji Bai 114
iCDPs (integrated conservation and
apes as relatives 16-17
in
201
iboga shrub 116
Society 28
of 144,
144, 145
de Recherche en Ecologie
Integrated Conservation and
Sierra
Humane
humans
murders
Nature
Institute for Tropical Forest
Liberia 368
Rwanda
Institut
apes 418,
seea(sopredation
within ape species, western
gorilla
117-18
Inter-African Forest Industries
Association IIFIAI 256-7

internally displaced
persons 368
IM
World Atlas
international
agreements and
conventions 246-50, 436
compliance with 249-50

International Fund for
Animal
langur208, 212, 214
Kenyah people 179
Lanjak-Entimau Wildlife Sanctuary.
Ketambe Research
Sumatra
151, 388,389,
Ketambe
The Last Great Ape 310
185, 186, 188,
international organizations 243-4
Ranger Federation
[IRF)436
initiatives
108, 110, 164,
Uganda
75,
7^,258,274,408,410,411
Ngogo community
79,
62, 63,
80
65
Primate Habituation Project
187
leaf-pile pulling
68
Leakey, Richard
12,
Islamic law 179, 198, 226, 227, 393
Leakey, Mary 12
leaves
in diet 31, 59, 98.
300,301,303,304
forest 30, 131-2, 340-1
eastern lowland gorilla

populations 140
Forest Reserve.
79
Union
326
enforcement 125
byapes44,
Burundi 304
international 246-50
45, 5S-?, 60-1,
national 245-6
71-2,73,85,90, 164, 187

of
humans
73, 144,
see also under individual
145,388,
389
Congo 325
countries
Kinabatangan Orangutan
Conservation Breeding
Group
95, 251
fi'edlL/s(91, 217-18,
240
West African Chimpanzee

Action Plan 251
Faunal Reserve. Congo 98.
legislation
killing
DRC
lUCN-Ifie World Conservation
Specialist
Lefini
kidnapping, of infant apes 65

kijang IMuntiacus muntjac] 172
Conservation Project 169-70,
regional agreements 250-1
Leguminosae
252. 437
Sabah 169-70,
181
lemurs 20
leopards 72, 112, 115, 135
Klaineanthus 107
Krokosua
38. 40. 69. 71. 107,
M9, 208
108,
Kinabatangan Wildlife Sanctuary,
Klainedoxa
clouded 195
Lesio-Louna Reserve 126, 326
1 1
Hills Forest
Reserve 356
Leuser Development Programme
Kutai National Park 181, 182
201,
202
252, 274
Leuser Ecosystem
422
37, 252,
Jane Goodall Foundation 326
fragmentation 199-201,420
Jane Goodall
history
Institutes 125, 224,
304,310,323,409,410,415,
John Aspinall Foundation 323, 326
Cameroon 125,305,311,
Julbernardia 40
317
Community Reserve 325
202, 252
lianas 173, 196

Liberia 227,
Lagoas de Cufada Natural Park,
132-3, ;58, 237, 257
conservation activities 149

populations 140, 141, 142
gorilla
tourism 150
Kalimantan
33, 161, 175, 180-1,
368
Guinea-Bissau 364
Lake Tanganyika Catchment
56, 367,
Reforestation and Education
legislation
182

life
land bridges 21
lifespan of
Kalinzu Forest Reserve 57-8
Landolphia
Likouala
Kanzi47-8
land reclamation, Indonesia 181
Karisoke Research Center 132, 133,

135, 138, 139,
149,389
and conservation
action 247, 368-70
Nature Reserve
Lana project 46
181,200
54,
370
Lake Victoria 410

Strict
apes
368
project ITACAREI 415
Lamandau
366-70
Ladia Galaska 202
Kahuzi-Biega National Park 60, 131,
201-2
Leuser International Foundation
lexigrams 46-8, 91
Lac Tele/Likouala-aux-Herbes
kabupaten202.i2]-2
200-2
legal recognition
Lac Lobeke National Park,
436
US
187.208
164.
use as tools 67
Kibira National Park, Burundi 78,
Isobertinia 40
244
Leakey, Louis 24-5
76-7
Ipomea 133
273
law enforcement 298
conservation success
invertebrates, as food 41, 60, 68-9,
community
Latin America,
Sumatra 194
Khoratpithecus 22-3
355,357
Ituri
River,
Malaysia 37, 255, 428
Laporfea spp. 133
Station,
;?2-3, 20;, 243, 252, 273
International Monetary Fund (IMFI
Itombwe
great apes 46-8, 91
Kelabit people 178
390,410,436
International
language IhumanI, acqusition by
178, 179
kebun 178
Kerinci-Seblat National Park 26-7
Welfare (IFAWI 436
Programme
Kayan people
08
Landsat images 223-5

land tenure. Borneo 178
history of
apes 217, 218
apes
88, 191,
218
swamps, Congo
124, 321.
323. 325
Limbe
Wildlife Centre.
272.312
Cameroon
Index
lions 73
Lufengpithecus 22-3
Lithocarpus spp. 164, 165
Lukuru
live
animal trade 229, 247, 345
Wildlife
91,95,338,339
bonobo 92
Luo Reserve
chinnpanzees 79, 247, 268-9,
for Scientific
action 247. 375-6
Research
374-5
Lwiro Primate Sanctuary 340
355, 359, 398, 402
and conservation
legislation
93-4
92,
376-7
Research Project
gorillas 143-4
Matlotus spp. 164
medical research 27-8, 370, 398
mandrill 344
orangutans 178, 179, 226

livelihoods, local
communities
280-2
Living Earth Foundation 310,
Lobelia spp.
437
16
lobeline 116
conservation
initiatives
258-60,
improvements 280-2
Africa 40, 49
maps, background data 10
Marahoue National Park, Cote
174,194,195
Marais, Eugene 24
MacKinnon, John 25
Marantaceae
Magnoliaceae 33
324
IMEMPI 415
Tanzania 44,
59, 60, 65, 66,
Maiko National Park
CAR 315
32,
populations
Maiombe
324
DRC 337-8
419
reducing 256-7
and adaptation
of
apes
Uganda 405
Bonobo Sanctuary
339-40
Lope National Park, Gabon

12i,
40, 60,
126,349,
gorilla studies 108,
Lopez, Eduardo 14
slow 187
Lossi Gorilla Sanctuary 119,1 20,
123,273,323,325,326
118
117,
Mayombe Forest 125

Mayombe massif 325
protected areas 249, 428
MbeliBai
Mbe Mountains Community
73-/4. 117, 123, 124,326
Forest
273
measles 147
chimpanzees
media 280-1
61
medicine, traditional 116. 359, 364,
413
perceived value of apes 323,
western 117
376
Bornean 166-7
Megaphrynium
108, 107, 109
macrostachyum 107
Sumatran 18-19, 190-1

subadult/subordinate 139, 153,
Meliaceae35, 107, 164, 185, 187
Mengame
Mali 371-7
Gorilla
Sanctuary 125,
305.310
menopause 88
menstrual cycle
371,373
56, 372,
182
123,321.322,
156-8
351
western
and conservation
337, 341
Mountains, Sierra Leone 397
63,
326
eastern 136-8. 139
Forest Reserve 85-6, 94,
79, 112, 121,
May,^NordBai
gorillas 99, 100
272,
181.
339
male behavior
logging companies 125, 256-7, 325
Borneo
Institute 91,
reserve,
Max Planck
427-8
191-4,256
Mawas
action 247, 428
Southeast Asia 37
gorillas 100
ecotourism 262
legislation
Indonesia 417, 419
61
orangutans 156-7
apes
428-30
impact on forests 224-5
maturation
425-7
181,222,273,
08,
Guinea 359
illegal 169, 177.
297
425
and gibbons 212-13
165 197
Gabon 348-51
09
fruiting 34, 38,
HO
forest 295,
Massif du Ziama 359
mast
Malaysia 425-30
Equatorial Guinea 121, 345
cordifolia
purpurea 109
Martyr, Debbie 26
335
concessions 40, 312
survival
131, 132,
Marantochloa
congensis 85
68,71, 79,413,415
Cameroon 121,307,312
38, 69, 85, 107, 108-9,
321,325
Mahale Mountains National Park,
and bushmeat hunting 256-7,
79,330
d'lvoire
MacArthur Foundation 303
Project
Borneo 168-70
loris,
Maninka people 376
long-tailed 174, 194, 195
Mahale Ecosystem Management
logging 220, 222-5, 255-7
Loma
mangroves 362, 364
ecology 173, 174,208
Sundaic 153
273-4,281-2,283,285,313
livelihood
Mangifera 35
mangosteens 178
87
14,
pig-tailed
communities 277
Lomako
golden-bellied 89
30;, 302
macaques
farming 374-5, 397
Lola ya
black 89
Mabanda/Nyanza-Lake. Burundi
disease transmission 330
Congo
mangabey
Mabali Scientific Reserve 337
livestock
local
373-4
apes
54,
bonobo
48.
88
chimpanzees 48-9. 62-3
a9
World Atlas
Africa 36, 39, 60. 301
gorillas 100
Southeast Asia 32. 33.35,
Meremia
INGASPI291,312. 326
national parks, see protected
175
79.
parks
228. 229. 342, 344, 345, 346
Nauclea ^08
Equatorial Guinea 74-5,
Mgahmga-Bwindi Impenetrable
Mont Peko National Park 328
254
Mont Sangbe National Park 328
Neesiaspp. 187,
Moraceae
nematodes,
Gorilla National Park.
Uganda
27. 131.
U6. 255.
261,408
35. 40. 58, 60, 133.
137
mosaic habitats 38-9.
MIKE (Monitoring
mother-infant bond
program
91,94
40, 50,
220
mountain
gorilla [Gorilla beringei
Cooperation (DGISI 351
diet 39, 98, 116
144,236-7,257,335,407
counting of 124. 141
Netherlands Development
coal 181
solitary 118
beringei]
bauxite 357
infanticide 138
nettles 133
diamond 396, 397-8
reproduction 139
Ngamba
gold 235
vocalization 142-3
coltan
impacts
of 49.
iron ore
359
24-5, 249, 387-9
Minkebe National Park
74. 123, 125,
405, 406
Minkebe-Odzala-Dja Interzone
Project 274
organizations
niche separation, primates 89-90.
218
analyses 19
gorilla
see ranging behavior
53.
17, 22, 24,
National Park 175. 178
97
Mount Kupe 311

Mount Nimba
56
molecular-clock analyses
19,
24
Nature Reserve
mouse deer
Mpassa Biosphere Reserve
swamp
70,
71-2
Congo Basin Wolfs 89
Musanga spp. 40,

Myhanthus 133
Diana 71
Myrtaceae
dryad 89
myths
cryptic
howler 24
leaf 31, 173-4, 194,
137.
58, 60
208
redtailed 59, 89, 90
vervet 61
montane
forests
57.
apes 54-5,
379
legislation
and conservation
380-1
[Pan troglodytes vellerosus]
consen/ation 309-12, 381-2
distribution 54-5, 57, 306, 307,
379, 380
apes 26-7
see also traditions and beliefs
threats to 307, 309, 380-1
Nini-Suhien National Park, Ghana
195,203
Old World 21
proboscis 172. 173
349. 352
Landscape, Burundi 301. 302
De Brazzas 89
forest 44
230-2
381-2
171, 172
Mukungu-Rukamabasi Protected
89
79,
action 79.247,311,312,
328-30, 359
monkeys
cercopithecine 41,
378
382
Strict
monkeypox 330
Allen's
Nigeria 378-383
Mount Kinabalu and Crocker Range
molecular analyses
Niger Delta 380
Cross River National Park 31,
389
Moaceae 87
chimpanzee
261.388
Program, Rwanda 147, 149.
orangutans 154
mobility,
?50,
Mountain Gorilla Veterinary
Mittermeier. Russell A. 244
318
407
tourism 149,
97
124. 315,
173-4. 194-5
threats to 143-8,386-7.405.
43.
53.56
CAR
populations 101, 140. 141. 151.
genetic variability 129, 148
DNA
NGOs. see nongovernmental

Ngotto forest,
mites, skin 147
chimpanzees and bonobo
see national great ape
habitat 132, 145-6
Miocene epoch 21-2
mitochondrial
NGASP
survival plan
distribution 129-31. 333, 336,
348, 349. 351
Chimpanzee
Island
Sanctuary 272. 410
77-8. 368
69, 71
nests
191
mining
bonobo 89
gorillas 100, 118, 141
orangutans 158, 167. 188, 190,
minerals, dietary 164
35, 164
chimpanzees
chimpanzees 65
Mimulopsis spp. 132, 133
192 196
nest building
Michael Leo Rion Sanctuary 312
Killing of Elephants!
190,
intestinal 79
Nephelium spp.
169-70. 187. 208

morbillivirus 235
of the Illegal
national
Ndegense people 93
Forest Conservation Trust
Mgahinga
named
areas and
Monte Alen National Park,
spp. 170
Messner. Reinhold 26
650
national great ape survival plan
355
Niokolo-Koba National Park,
Senegal391.392, 393, 394
Nabolongo island 315
Nishida, Toshisada 244
National Geographic 25
Nomascus, see gibbons, crested
Index
nongovernmental organizations
diet
INGOsI 243, 251-2
and foraging 34-5.
|PanEco-SOCPl437
social 156-8
Africa251-2, 310-12, 339
pangolin, tree 44, 60
Congo 326
conflict with people
DRC 339
differences between sexes 154.
fund raising 274
Norwegian Agency
for
distribution32. 33. 35-6. 418.
298
evolution 21, 22-3. 153-4

Parl<,
122,321,
324, 326
346
fiigtilands
Nyungwe
National Park, Rv\/anda
276, 387-8
Pan troglodytes
populations 218
parasites 79. 147
research 25, 159, 182
parasol tree 60
sanctuaries 267, 270
Par/nan spp. 58
taxonomy
park rangers
13,
153-4
murder
role
tourism 264-5
key
Sumatran orangutan
Organisation pour
79, 107, 120, 122, 123,
321,324-5,326
palm
60, 71,
products 2S0-
parrot. African grey
lODEBI 303
partnerships 242-5
trade
in
325
see also transfrontier
conservation
268-9, 359
Angola 295. 298
western gorillas 126
see a/so sanctuaries
party formation
bonobo 85-6
Ouranopithecus 23
okapi 44
see a(so groups
Outamba-Kilimi National Park 397,
Okapi Faunal Reserve 237. 335
okoume l4ucoumea klaineana] 349

Okwangwo-Takamanda
Paspa/um
399
Owen, Richard
16,
peatlands, distribution
peat-swamp
monkeys
Omomycidea 19
project 151
Sylveslri',
Anatomy of a Pygmie
compared with that of a
or the
'Monl<ey',
palm
oil
plantations, see
oil
drainage
palm,
see also
plantations
Orangoutang, sive 'Homo
an
'Ape'
forests 174, 175,
disturbance 158-9, 168-9
Pa//sofal09, 133
16-17
Initiative'
and a
'Man' ITysonl 15
palms 164
181-2
177,
swamp
Per/p/us (Hannol
climbing 33
Pan African Sanctuary Alliance

IPASAl 267, 272, 340, 398,
Petit
14,
pet trade 345
437
chimpanzees
Panbanisha 47-8
402, 407
79, 268-9, 355,
Pancovia laurentii 87
Pandanaceae 109
Peucedanum linden
Pandanus
Philippines,
Orangutan Foundation 182. 243.
spp. 109
PanEco Foundation
for Sustainable
Development and
orangutans
Intercultural
betiavior
orangutans 178, 179, 198,226
Exchange & the
Sumatran Orangutan
'\33
community
initiatives
273
Pandrillus272, 312, 382
Orangutan Network 252
16
Loango region 116
Orangutan Conservation Forum

252. 423
forests
Penan peoples 174
orang pendek 26-7
252. 423. 437
Southeast
176-7, 196
21
the Origin of Species (Dar/in|
'Open
Asia 176-7
area 382
On
spp. 170
paternalism, bonobo 87
17
transboundary protected
Old World
345
logging companies 125, 256-7,
74-5. 78, 79, 267,
orangutan 179
Gabon 348
10
named partes
Defense de
268-9
production 49, 77-8, 320
maps
I'Environnement au Burundi
chimpanzees
427, 429
oil
la
to
see also protected areas and
orphaned apes 245. 246
158
plantations 158. 181, 194,417,
144. 145
148
parks
see also Bornean orangutan;
oil
verus, see western
chimpanzee
habitat 35, 37
240,419-20
Congo
troglodytes, see
central chimpanzee
threats to 158-9. 169-70,226,
Odzala-Koukoua National Park,
chimpanzee
eastern chimpanzee
early description 15-16
108. 109,
troglodytes, see
Pan troglodytes marungensis 53

Pan troglodytes schweinfurthii, see
Pan troglodytes
preindustrial 155-6
Cooperation INORADI 297.
Congo
Pan
419
Development
Nouabale-Ndoki National
Nsork
154. 155
named NGOs
Panicum brevifoUum 85
Pan paniscus, see bonobo
Panpanzee 47-8
differences between species
development projects 259-60
see a(so
258
254
integrated conservation and
Programme
Conservation
154-5. 163-4
pig
bearded 163, 165 168. 172

Sundaic 153
Pigafetta. Philip 14
451
World Atlas
apes
135
Pilbeam. David 22
of great
plantations, forest conversion 158.
see also bushmeat hunting;
429
181, 19A, 397, A27,
15, 115,
transfrontier 125, 245, 255, 305,
310-11,317,382
Uganda
hunting
pregnancy, age at
plants
first 88,
27, 79,131,
146,247,
249,255,261,406,408-9
218
260
evolution 20
prey species 44
'protein projects' 259,
gibbon diet 208
primates
Proyek Lahan Gambut 181
important food species 38, 60
evolutionary origins 19-21
Pterocarpus tinctonus 71
leguminous38,
niche separation 89-90. 173-4,
public education 123. 264, 271-2,
40, 71, 108,
282-3,285.311
194-5
M9-70
mountain
private sector 125, 256-7, 277.
gorilla diet 133
35
283
Soutfieast Asia 33
proboscis
w/estern gorilla diet 108-10
Proconsul
orangutan
diet
Purchas. Samuel 14-15
unique features 18
nnedicinal value 7!^
monkey
172.
mountain gorillas
western
M3
see a/so bushmeat hunting;
bonobos 93-5
hunting
Podocarpaceae 39
RamsarSites303, 325, 364
'bottom up' 273
Ranalisma humile 85
Burundi 300.303
range states 288-9. 438
Cameroon306,
unrest 233-5, 236
308, 309, 311,
Congo 321
CAR
DRC94, 332
chimpanzees
316, 317-19
79. 80,
historical conflict
369, 375.
DRC
see Sumatran
334, 335, 336, 337, 338-9
effectiveness of 79,80, 123,
orangutan
Pongo pygmaeus mono 6 180

Pongo pygmaeus pygmaeus 161,
1
chimpanzees 44-5, 61-2
Gabon340,
gibbons 209
Ghana
348, 34?, 350, 351,
gorillas 98, 99
eastern lowland 98, 134
354, 355-6
mountain
gorillas
180
Pongo pygmaeus wurmbii
161,
79
population and habitat viability
assessment IPHVAI 251
mountain 387-8, 408-9,
124. 141
decline
Bornean orangutan
180, 787
98,
Bornean 164, 166

Sumatran 187-8
western 123, 125
Guinea-Bissau 363, 364
Raphia spp. 107, 109
Indonesia 418, 420-2
rattans 33
l<ey to
maps
/?edl/s7(IUCNI 91, 217-18, 240
10
projected 240
Liberia 367, 369
reforestation 224
rate of 219
Malaysia 418, 428
refugees,
estimated numbers 218, 219

viability in
protected areas 253
porcupine 172
brush-tailed 229
predation Iby great apesi 41, 44, 187
in
existing
rangers 148
Rwanda
144-5, 146, 332,
regional agreements 250-1,
opportunity costs 254
populations
human
335, 368
Mali 372, 375
role of
134
western 111-12
orangutans 155, 156
410
Guinea 357, 359
populations
see also individual countries

ranging behavior 141
Equatorial Guinea 343, 345
352
of international
bonobo 90
125,254
orangutan
Pongo pygmaeus, see Bornean
Index
conventions 246, 247
Cote d'lvoire 328-30
Polygalaceae 35
Human Development
membership
Congo 322, 324-5
353, 355
Guinea-Bissau 362
219-20
233-5
rankings 230
403, 408-9
Gabon 348
EIA legislation 245-6

forest cover decline
312
Angola 295
censuses
282.287.311
Angola 296-7
abelii.
Quercus spp. 165
protected areas 230-2, 240, 252-5.
alternatives to 260-1
Pongo
Q
quarantine sites 421, 422
126,352
poaching 144, 386
Ghana
de
Projet Protection des Gorilles IPPG)
gorilla 7/4
PoaceaeBS. 133
political
et
IPERCSI 393-4
pneumonia U7, 235-6, 386-7
76
Chimpanzes du Senegal
gibbons 210
pythons 168
173
Recensement des
chimpanzees 68
Pycreus vanderysti85
pygmy peoples
Programme d'Education
play
452
Tanzania 414, 415
predators
pigeons 171
384, 387-8
bonobo 45
Senegal 392, 393
chimpanzees 58
Sudan 401. 403
23-4
273-4
reintroductions, see release of
captive apes
release of captive apes 266-7,
270-1
Index
bonobo 339-iO
chimpanzees 74-5.
threats to great apes 750, 220.
386-7
79, 253,
267. 268-9.2TI. 326.356.
tourism
habitat monitoring 151

150, 261,
388
Savage,
370,^13.415
Thomas
16
Savage-Rumbaugh. Sue 46-7
213-U
gibbons
413
forest losses 225-5,
savanna woodlands
40. 45. 58. 107
gorillas 126, 271
Schaller.
orangutans 180, 264. 267, 270,
Sebangau National Park
271,422
Saba
remote sensing
Renealmia africana 85
secondary forests
425
88-9
bonobos 87
degraded forests 169-70
chimpanzees
chimpanzees 48-9,
68
apes 161,
425,426,427
eastern gorillas 139, 141
40. 107.
40, 70
gorillas 40. 112. 135
mechanisms
171
gorillas 100
great ape habitat 32. 35
non-ape fauna
threats to great apes 427-8
orangutans 170-2. 194
218
sacred
reproductive interval 118,218
bonobo
95,
218
88,
Guinea 359
gorilla 118, 119
57,
influential
programs 272-3
Senecio
sp. 116.
132
Africa 267
391
Cameroon 312
58-9, 66
Sendje. Equatorial Guinea 228
Senegal 391-4
sanctuaries 242. 243, 266-7
chimpanzees/bonobo 50-1,
chimpanzees 370. 382. 398. 410
Congo 323, 326
great ape habitat 58
see also
DRC 339-40
legislation
studies
reserves, see protected areas
and
named reserves
educational value 272
apes
54,
56.391.392
orangutans 159, 182,202-3

field
264
rehabilitation site
268-9
see bais
Center 311
captive great apes 25-8
Semenggoh
Sendje (orphaned chimpanzee!
sambar deer 172

Sanaga-Yong Chimpanzee Rescue
research
171. 172
seed predation 171-2. 187
337
salt clearings,
western
sites.
Salonga National Park 91, 92, 93,
reproductive rate
169-70
seed dispersal

65.
175.
175
conservation 182,428-9
49,
132
24. 27.
176-7. 181
bonobo
59-60
Sabah
413
151. 223-5,
florida
George
and conservation
action 247, 393
Gabon 352
Ghana 356
rhinoceros 171, 194
Rhynchospora spp. 109
Liberia 370
World Heritage
Rio
Campo
Reserve, Equatorial
rehabilitation
Guinea 346
202 238-40.
338,
132, 133
Rubiaceae39,
Rubondo
108, 133, 170.
208
Island, Tanzania 253, 271.
Sierra Leone 398
sexual behavior
bonobos 48-9. 86-7
gorilla 126
San Diego Zoo
17,
chimpanzees 48-9. 62-3
310
gorillas 100
Rumonge
Sapo National Park,
185, 187
35, 87.
homosexuality 137. 191
164
non-reproductive 88
Liberia 369
Sapo people 227
301
Sapotaceae

war/genocide 144, 146,
Sarawak
gorillas 100. 139
apes
56,
384, 385-6
389-90
and conservation
apes 425-7
great ape habitat 32, 34, 35
Sarcophrynium spp. 107
sassy tree 69, 71

satellite data
Sberman 46-7
shrews, elephant 44. 61
schweinfurthianum 85
action 247. 276, 387-9
180,
425
conservation 182, 428
234, 332, 335, 383, 385
sexual development
bonobo 88
85, 108, 164
383, 385
legislation
254
Uganda 410
western
Sapindaceae
Rwanda 383-90
427
sex ratios, at birth 117. 191
beccahanum
Forest Reserve, Burundi
249
Sandoricum 196
413,415
Site
Sepilok rehabilitation project 264.
sex differences, orangutans 154.
Rubus spp. 133

Rumbaugh, Duane 46-7
civil
of
370
420
Rosaceae39,
and release
apes266-7, 270-1,326. 356,
Rio Muni 342-4
road building 126,
393
siamang ISymphalangus] 194-5,

206.208-11.214
Sierra Leone 395-9

395. 397
56, 396,
apes
54,
397
1(53
World Atlas
field
398-9
and conservation
death of
sticks,
Ul
]38.
gorilla
73,
74,
115,
Paul 7A
Sivapithecus 22, 23
U7
skulls, trade
in
U8,
260
229, 317, 323, 338,
119,318
gorillas 102, 107, 109,
775,
females 62-3
'swollen'
orangutans 153, 156-8, 188-9
sympatry, chimpanzees/gorillas
135, 757
Symphalangus. seesiamang
Syzygium 137
400
apes
55, 56,
401
403-4
chimpanzees 45-9,
5?, 62-5,
legislation
and conservation
74,
115,
117-18

soils, eating of 60, 133,
164
songs, gibbon 210-11
Southeast Asia
Programme ISOCPI
Leone 398, 399
Sierra
Tai National Park, Cote d'lvoire 61

62, 65, 68, 72, 328,
252, 273,
330
conservation success 79
abetiil

diet
see also traditions and beliefs
Tacugama Chimpanzee Sanctuary,
33-7
421,422-3,435
192-3
198,226,227
330, 393
bushmeat 226
Sumatran Orangutan Conservation
infanticide 138
orangutans 156-8
16
taboos 94, 323, 330, 338, 344
Sumatra
biogeography and ecology 32,
eastern 135-9
western
Islamic law 179,
402
gorillas 99-101, 115, 117
Tabernanthe iboga
Tabin Wildlife Reserve, Sabah 174
action 2^7. 402-3
threats to great apes 220. 400,
349, 351
gibbons 209-11
Tamabu Range 175

Tanganyika Catchment,
Reforestation and Education
and foraging 185, 187,
Project 224
biogeography 32, 33
196-7
ecology 33-7
ecological role 194
ecotounsm 262
interactions 194-5
167, 168, 181, 182, 262,
great ape distribution 287
nest building 190
423
infrastructure development
238-40
NG0s252
tsunami (December 20041
ranging 187-8
tannins 59. 164
tantalum 236, 257
named
countries
Spatholobus spp. 170

species action plans 250-1
202-3
tapirs 194
orangutan 154, 155

distribution 185, 186,418,
eastern chimpanzee 410-11
habitat 195-7
391,394,398
Borneo
33,
34
335-7
Tchimpounga Chimpanzee
278
area needed for
200
ITGHKl 420-1
taxonomy 13
Biology ITCCBI 437
populations199, 217, 27S, 219
species richness, Sumatra and
Cuna Hutan Kesepakatan
Tayna Gorilla Reserve 131,
disturbance 191-4, 199-202

lifespan
Tata
Tayna Centre for Conservation
419
evolution 153-4
species data 10
Tanzania
bonobo 251
western chimpanzee 356, 377,
2,
Tanzania see United Republic of
use 189-90
differences from Bornean
201-2
Tanjung Puting National Park 164,
social behavior 188-9, 192-3

tool
sanctuaries 267, 270
see also
775-7,
196
bonobo i5-9, 85-8
654
187,
orangutan 164, 174,
74, 400,
3AA, 386
social behavior
and
168-9, 777, 181-2
Suaq Balimbing, Sumatra
227
snails, African giant 259,

78,
clearance and drainage 158-9,
Sudan 400-4
226,
forests 38-9
bonobo 85
subadults
simians, origin 20-1
bearded
chimpanzees 318, 321, 323
strategic conservation priorities
192-3
117
snares
swamp
67. 68, 69, 190,
282-3
136-7
pig,
sustainable development 259-60
193
192.
eastern gorillas 97, 134, 135,
mites
use as tools
168, 171, 173
Sundaland 153-4
Su5 barbatus. see
126, 351
Sterculiaceae U6. 770, 187
communication 1^3
sl<in
Chimpanzes
stepping sticks' 68
silverbacks 97, 99
Sita,
sunbears
giant 168
apes 220,
397-8
western
threats to 198,218
60
Station d'Etudes des Gorilles et
action 2A7, 398

tfireals to great
190-1,278
squirrels 69, 72, 173

flying 44,
legislation
sport hunting 179
studies 68
viability 197,
Sanctuary 326
teeth, fossil
television
remains 23
280-7
740,
Index
termite-dipping 67, 68
loss of 179
termites 60, 61, 111, 164
medicinal/health value
home range
see
territory,
407-8
legislation
of
apes
threats to great apes 220. 405,
323, 376
407
Tetrameristaceae 183. 196
medicinal plants 116
Tetrapleura spp. 108
Uganda
Wildlife Authority 261
Thelypteridaceae 85
sacred sites 359
Uganda
Wildlife Education Centre
Threat Reduction Assessment ITRA)
(ua species 178,
415
tourism
227
283, 299
Cameroon 125,305,311,317
status classification 217-18
CAR 317
apes
182.
217-19
see also
tiger,
threats
and
Mali 374, 377
United Kingdom (UKl 231, 369
Nigeria 382
Tiwai Island Wildlife Sanctuary 397
United Nations Development
translocation of great apes 270-1,
United Nations Educational,

Scientific
304
tool use 17
travel,
chimpanzees
Treculia afncana
67, 68-9,
361
gorillas 100-1
TrichiUa spp. 107
Trinationale de
192-3
Programme lUNEPl
see ranging behavior
Sumatran orangutan 189-90.
la
GLOBIO
Sangha
Agriculture Organization of
apes as 143
CAR 318
Tropical Forest Trust 257

76-7, 330, 361
the|FAOI219, 245, 386

United Nations Foundation 94, 274
Tropical Rainforest Heritage of
United Nations Mission
Sumatra World Heritage
235-6, 262-3, 264, 407,
Site
422
costs/risks of 123, 126, 147,
Liberia
United Republic of Tanzania 412-16
tsunami. Southeast Asia 201-2
412-13
Cote divoire 330
Tulpius. Nicholas 15
DRC 339
Tu5i(437
Tutm. Caroline
gorillas 123, 126, 149, 750,261,
twigs, use as tools 193
388,410
in
366
(uaspeciesl78, 227
410
computer model
United Nations, Food and
trophies,
Congo 325
236-7, 239
125, 305.
Thumfetta 133
benefits of 149, ;50, 262. 410
chimpanzees
243-4,
290
87
311.317.325
tourism 242, 261-3
94.
344, 386
89, 91
(UNESCOI
151,243-4,245,248,290,403
148, 22S, 229, 317, 323, 338,
bonobol7,
and Cultural
Organization
trapping/snaring of great apes 78,
288
150.
Programme lUNDPl 297-8
TransGabonais railway 348-9
Toepfer, Klaus 275, 433

56,
WCMCI437
Senegal 393
timber extraction, see logging
Togo
Monitoring Centre lUNEP-
Guinea 255
species
Sumatran 195
spp. 770
UN Global Compact 257

UNEP World Conservation
Ghana 356
under each great ape
Tiliaceae 110, 133
Umbelliferae 133
Uncaha
Congo 125,325
Gabon 125
named
Ulmaceae 164
Ulu Kinabatangan, Sabah 182
muttiple 237-8
vulnerability of
75-7,261-2,410
272,410
transfrontier conservation 125, 255,
50,
chimpanzee releases iRubondo

islandl253, 271,413, 415
74.
126
chimpanzee studies 63
twins 65
apes
55,
57,413,414
habituation of apes 76-7
orangutan 264-5
Rwanda 750,261,388
Uganda 75-7,261-2,410
western lowland
gorilla
318
416
legislation
bacl<ground and
trade
bushmeat 226-9, 245

forest products 224
live
apes
79, 178, 179, 198,226,
229, 247, 268-9, 355

traditions
230,
Urera 133
Urticaceae 133, 187
Park 244, 261-2

conservation actions and
protected areas
79, 247, 249,
226, 227,
323, 338, 344-5
chimpanzees 330, 359, 376
Urtica massaica
US Agency
33
for International
Development (USAID)
95.
245,274.318,351,403,
apes 405,
415
406
410-11
413, 415
413
Bwindi Impenetrable National
255.261,408-9
and beliefs
bonobo 92-3, 94
bushmeat hunting
economy
247
404-5
Borneo 179
and conservation
action 7?,
Uganda 404-11
US
Great Ape conservation Act

120001 274
455
World Atlas
fvlali
V
386.
people 227
DRC98.
impact
102, 130.
fiuman
of
western
131.335.340
conflict 144-5.
response
gorilla tiabitat 132.
295, 438
World Heritage Sites 123. 248. 249.
112. 115
to
338. 359. 422
disturbance 118
gorilla populations
habitat 99. 105. 107
chimpanzees 68
lifespan
290
76
World Zoo and Aquarium

Conservation Strategy 263
World Zoo Conservation Strategy
218
263
gibbons 210-11
Sumatran orangutan
gorilla
189. T)2
148.388.389
Organization 151. 230-2. 243.
251-2.310.331.438
research 125-6
field
Karisoke Research Center 132.

133. 135. 738.
censuses 724
reproduction 119
19
Volcanoes National Park. Rwanda

41. 131. 134.
Wrangham. Richard 27
VWVF-The Global Conservation
population 119-20. 218
742-3
gorilla
gaps
studies
73-74
knowledge 103
in
observational challenges 105
149.389
Volcanoes Safaris 437

vulnerability of
distribution 102. 105. 106
9'i
threats to 120-3. 126
apes 182. 217-19
western lowland
307.302
gorilla [Gorilla
35. 164
Xanthopyllaceae 35. 164

Xylopia 107
gorilla gorilla]
distribution 294. 295. 305.
X
Xanthophyllum spp.
tourism 123. 126
Vyanda Forest Reserve. Burundi
310
Development. Johannesburg
medicinal value of foods
bonDbo89.
institute
World Summit on Sustainable
conservation 123-5
diet 98-9. 107-11. 773
740, 141
vocal betiavior
mountain
World Heritage Species 248-9

World Resources
social 774. 115. 117-18
133
mountain
308
Cameroon 315. 316

Congo 321. 322
DRC
333. 335. 336
Eguatorial Guinea 342-4
Wallace, A.R. 16-17
Gabon
V\/amba region 88
warfare, see
World Heritage Convention 248,
communication 119
nest building 118
Virunga massif
western
lUCN-The World
Conservation Union
behavior
chimpanzees
Rwanda 388-9
mountain
World Conservation Union see
gorilla [Gorilla gorilla]
interactions with
146.234
318, 325, 357,
369
374-5. 397-8
deforestation 388-9
27
Rev Mr 16
World Bank 246, 257,
Wilson.
threats to 330. 359. 364. 368.
Virunga National Park 139
Forest 388
409, 410
Williams, Shelly
releases of 15.253.271.413
389
human
Yaounde Declaration 250.311.345.
348. 350
threats to 101-2.323-4
conflict
water, use of/access to 57. 60.
74
wealth, increasing local 280-2
351
yaws
wetlands
distribution
in
Southeast Asia
176-7
W/ehdjeh clan 368
West African Chimpanzees: Status

Survey and Conservation
Action Plan 356. 377. 291.
see also
Yeti'
123.
324
26
Yushania alpina [Arundinaria alpina,

bais:
forests;
bamboo|39,
peat-swamp
swamp
133. 134
forests
WM Chimpanzee Foundation IWCFl

271-2.330-1.370.437
394. 398
western chimpanzee [Pan

troglodytes verus]
Unit. Oxford
Research
346
Wildlife Consen/ation Society
behavior 62
conservation actions 368-70.
Zaire,
IWCSI
243. 256. 339. 438
see Democratic Republic of

the
Congo IDRCI
Zambia 289
Burundi 304
Zemongo Faunal Reserve 317

Zingiberaceae38. 85, 107, 164
Cote d'lvoire 328-30
Cameroon 310. 311

Congo 324. 325
Ghana 354, 355
eastern gorilla states 142. 151
Zoological Society of London 346.
Guinea 357-9
Gabon 351-2
Guinea-Bissau 362-4
Nigeria 381
Zoological Society of Ivjilwaukee 95
Project for the Conservation of
Zoological Society of San Diego 310
398-9
distribution 54-5, 56
Liberia 367,
656
Uganda
Sierra Leone 396. 397
U9.
veterinary assistance 147,
Vili
Nyungwe
372, 373-4
Senegal 391, 392
368
200-based conservation 263-6

398, 438
JULIAN CALDECPTT
He has
an ecologist and primatologist by
js
studied gibbons and
macaques
ests and has worl<ed on wildlife
in
Malaysian
management and
training.
rain for-
biodiversity
conservation throughout the global tropics, including projects

to conserve
Bornean and Sumatran orangutans, western and
Nigeria-Cameroon chimpanzees, and Cross River
lERA MILE'
is
a biologist with the
Monitoring Centre
who
UNEP World
gorillas.
Conservation
analyzes threats to biodiversity rang-
ing from the global to a local scale.
Her specialties include
species mapping and modeling, vulnerability assessment.
and
priority setting.
Copub:
UNEP-WCMC
Jacket design by Victoria Kuskowski

Jacket
illustrations: Front:
Powles/Still Pictures;
{from top
left);
chimpanzee {Pan
troglodytes),
Bornean orangutan (Pongo pygmaeus),
Pictures; western gorilla {Gorilla gorilla),
T. J.
Rich/naturepl.com;
bonobo {Pan
paniscus), Cyril Ruoso/Still Pictures; Sumatran orangutan {Pongo
Anup Shah/naturepl.com;
abelii),
eastern gorilla {Gorilla beringei), Martin Harvey/Still
Pictures. Back: (top to bottom):

Still
Mike
Cyril Ruoso/Still
chimpanzee {Pan
Pictures; eastern gorilla {Gorilla beringei),
troglodytes),
Mike Powles/
Bruce Davidson/naturepl.com.
IN
ASSOCIATION WITH THE UNITED NATIONS GREAT APES SURVIVAL PROJECT (GRASP)
"Great apes are self-aware, complex communicators, and skilled exploiters of their environment. Yet the
most self-aware, the most complex com-
municator, the most exploitative of

extinction. This
living relatives
all, is
now threatening
and
lays a basis for
how we can
and
COLIN
GROVES, author of Primate Taxonomy
and analysis relevant to conservation
and extent
ALEXANDER
of material
H.
preserve them for our
theirs."
"A well-researched, up-to-date, clearly presented

tion
the others with
book records what we humans know about our closest
posterity
P.
.#r
compendium
of informa-
of the great apes. This
amount
has never before been brought together."
HARCOURT, university of
California, Davis
Provides the most up-to-date and
comprehensive data available
Contains more than
200
full-color
photos
Includes nearly
and diagrams
Printed in the United
Kingdom
UNIVERSITY OF CALIFORNIA PRESS Berkeley
94704 www.ucpress.edu
50
full-color
maps

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World Atlas of Great Apes and Their Conservation

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WORLD ATLAS OF

ESPITE THE DEDICATED EFFORTS OF

and organizations, the great apes our closest

of extinction. This sw/eeping atlas provides

a comprehensive overview of what

apeschimpanzee, bonobo, Sumatran

orangutan, Bornean orangutan, eastern

association with the United Nations Great

background on behavior and ecology

book gives a thorough

ing detailed habitat requirements, the apes' ecological role,

and the possible consequences

rent conservation efforts,

each species across

description of the threats, cur-

and additional protection needed

Many full-color maps and

a broad readership, from specialists and policymakers to general

readers concerned about the survival of these charismatic

This book represents the work of a dynamic alliance of

of the world's leading great

ape research and conser-

vation organizations. Bringing together United Nations agen-

governments, foundations, and private-sector interests,

the project aims to raise the international profile of great ape

conservation and to build the

political will for further action.

being done by specific organiza-

support of great ape conservation and where conser-

most needed and most

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

University of California Press

Berkeley and Los Angeles, California

Cambridge. CB3 0DL.

+a 101 1223 2773U

book may be reproduced by any means, or

transmitted or translated into a machine language witfiout

volume do not necessarily

contributory organizations, editors, or

publishers. The designations employed and the presentations do

UNEP-WCIvIC or contributory organizations,

concerning the legal status of any country, territory,

authority, or concerning the delimitation of

boundaries, or the designation of

Clothbound edition ISBN: 0-520-24633-0

the Library of Congress.

eds 120051 World Atlas of Great Apes

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

Great ape distribution

UNIVERSITY OF CALIFORNIA PRESS

of Great Apes and their Conservation

and their Conservation

219 Huntingdon Road

Cambridge CB3 DDL. UK

the principal United

sets the global environmental agenda,

promotes the coherent implementation

within the United Nations system, and that serves as an

authoritative advocate for the global environment.

Lee Shan Khee

objectives include analysis o* the state of the global

environment and assessment

environmental trends, provision

early warning information on environmental threats,